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Birds are a group of
warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The onl ...
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s constituting the
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
Aves (), characterised by
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s, toothless beaked jaws, the laying of hard-shelled eggs, a high
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
rate, a four-chambered
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
, and a strong yet lightweight
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
. Birds live worldwide and range in size from the
bee hummingbird The bee hummingbird, zunzuncito or Helena hummingbird (''Mellisuga helenae'') is a species of hummingbird, native to the island of Cuba in the Caribbean. It is the world's smallest bird. Description The bee hummingbird is the smallest living ...
to the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
. There are about ten thousand living species, more than half of which are
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
, or "perching" birds. Birds have whose development varies according to species; the only known groups without wings are the extinct
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
and
elephant bird Elephant birds are members of the extinct ratite family Aepyornithidae, made up of flightless birds that once lived on the island of Madagascar. They are thought to have become extinct around 1000-1200 CE, probably as a result of human activity. ...
s. Wings, which are modified
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
s, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including
ratite A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites. The systematics ...
s,
penguin Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s, and diverse
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s and some
waterbirds A water bird, alternatively waterbird or aquatic bird, is a bird that lives on or around water. In some definitions, the term ''water bird'' is especially applied to birds in freshwater ecosystems, although others make no distinction from seabi ...
, have further evolved for swimming. Birds are feathered
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and constitute the only known living dinosaurs. Likewise, birds are considered
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s in the modern
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
sense of the term, and their closest living relatives are the
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
ns. Birds are descendants of the primitive
avialans Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally use ...
(whose members include ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'') which first appeared about 160 million years ago (mya) in China. According to DNA evidence, modern birds (Neornithes) evolved in the Middle to
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
, and diversified dramatically around the time of the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
66 mya, which killed off the
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s and all non-avian dinosaurs. Many
social species Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
pass on knowledge across generations, which is considered a form of culture. Birds are social, communicating with visual signals, calls, and
songs A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetition ...
, and participating in such behaviours as
cooperative breeding Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers. Cooperative breeding encompasses a wide variety of group ...
and hunting, flocking, and
mobbing Mobbing, as a sociological term, means bullying of an individual by a group, in any context, such as a family, peer group, school, workplace, neighborhood, community, or online. When it occurs as physical and emotional abuse in the workplace, suc ...
of predators. The vast majority of bird species are socially (but not necessarily sexually)
monogamous Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
, usually for one breeding season at a time, sometimes for years, and rarely for life. Other species have breeding systems that are
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
(one male with many females) or, rarely,
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
(one female with many males). Birds produce offspring by laying eggs which are fertilised through
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
. They are usually laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching. Many species of birds are economically important as food for human consumption and raw material in manufacturing, with
domesticated Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
and
undomesticated Wildlife refers to undomesticated animal species, but has come to include all organisms that grow or live wild in an area without being introduced by humans. Wildlife was also synonymous to game: those birds and mammals that were hunted fo ...
birds being important sources of eggs, meat, and feathers.
Songbird A songbird is a bird belonging to the suborder Passeri of the perching birds (Passeriformes). Another name that is sometimes seen as the scientific or vernacular name is Oscines, from Latin ''oscen'', "songbird". The Passeriformes contains 500 ...
s, parrots, and other species are popular as pets.
Guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
(bird excrement) is harvested for use as a fertiliser. Birds figure throughout human culture. About 120 to 130 species have become
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
due to human activity since the 17th century, and hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them. Recreational
birdwatching Birdwatching, or birding, is the observing of birds, either as a recreational activity or as a form of citizen science. A birdwatcher may observe by using their naked eye, by using a visual enhancement device like binoculars or a telescope, b ...
is an important part of the
ecotourism Ecotourism is a form of tourism involving responsible travel (using sustainable transport) to natural areas, conserving the environment, and improving the well-being of the local people. Its purpose may be to educate the traveler, to provide funds ...
industry.


Evolution and classification

The first
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of birds was developed by
Francis Willughby Francis Willughby (sometimes spelt Willoughby, la, Franciscus Willughbeius) FRS (22 November 1635 – 3 July 1672) was an English ornithologist and ichthyologist, and an early student of linguistics and games. He was born and raised at M ...
and
John Ray John Ray FRS (29 November 1627 – 17 January 1705) was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after ...
in their 1676 volume ''Ornithologiae''.
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
modified that work in 1758 to devise the
taxonomic classification In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
system currently in use. Birds are categorised as the biological class Aves in
Linnaean taxonomy Linnaean taxonomy can mean either of two related concepts: # The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus t ...
.
Phylogenetic taxonomy Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
places Aves in the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Theropoda Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ca ...
.


Definition

Aves and a sister group, the order
Crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
, contain the only living representatives of the reptile clade Archosauria. During the late 1990s, Aves was most commonly defined
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
as all descendants of the
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of modern birds and ''
Archaeopteryx lithographica ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
''. However, an earlier definition proposed by
Jacques Gauthier Jacques Armand Gauthier (born June 7, 1948 in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology. Life and career Gauthier is the so ...
gained wide currency in the 21st century, and is used by many scientists including adherents to the ''
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
''. Gauthier defined Aves to include only the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of the set of modern birds. This was done by excluding most groups known only from
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
, and assigning them, instead, to the broader group Avialae, in part to avoid the uncertainties about the placement of ''Archaeopteryx'' in relation to animals traditionally thought of as theropod dinosaurs. Gauthier and de Queiroz identified four different definitions for the same biological name "Aves", which is a problem. The authors proposed to reserve the term Aves only for the crown group consisting of the last common ancestor of all living birds and all of its descendants, which corresponds to meaning number 4 below. He assigned other names to the other groups. # Aves can mean all
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s closer to birds than to
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s (alternately
Avemetatarsalia Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinos ...
) # Aves can mean those advanced archosaurs with feathers (alternately
Avifilopluma Avifilopluma ("bird filoplumes") is a clade containing all animals with feathers. Unlike most clades, which are defined based on relative relationships, Avifilopluma is defined based on an '' apomorphy'', that is, a unique physical characterist ...
) # Aves can mean those feathered dinosaurs that fly (alternately
Avialae Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used ...
) # Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
", in this sense synonymous with Neornithes) Under the fourth definition ''Archaeopteryx'', traditionally considered one of the earliest members of Aves, is removed from this group, becoming a non-avian dinosaur instead. These proposals have been adopted by many researchers in the field of palaeontology and bird evolution, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is often used synonymously with the vernacular term "bird" by these researchers. Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s closer to birds than to ''
Deinonychus ''Deinonychus'' ( ; ) is a genus of dromaeosaurid theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period, about 115–108 million ye ...
''", with ''
Troodon ''Troodon'' ( ; ''Troödon'' in older sources) is a wastebasket taxon and a dubious genus of relatively small, bird-like dinosaurs known definitively from the Campanian age of the Late Cretaceous period (about 77  mya). It includes at least ...
'' being sometimes added as a second external specifier in case it is closer to birds than to ''Deinonychus''. Avialae is also occasionally defined as an apomorphy-based clade (that is, one based on physical characteristics).
Jacques Gauthier Jacques Armand Gauthier (born June 7, 1948 in New York City) is an American vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology. Life and career Gauthier is the so ...
, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered wings used in flapping
flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...
, and the birds that descended from them. Despite being currently one of the most widely used, the crown-group definition of Aves has been criticised by some researchers. Lee and Spencer (1997) argued that, contrary to what Gauthier defended, this definition would not increase the stability of the clade and the exact content of Aves will always be uncertain because any defined clade (either crown or not) will have few synapomorphies distinguishing it from its closest relatives. Their alternative definition is synonymous to Avifilopluma.


Dinosaurs and the origin of birds

Based on fossil and biological evidence, most scientists accept that birds are a specialised subgroup of
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and, more specifically, members of
Maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
, a group of theropods which includes
dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
and
oviraptorosaurs Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or ...
, among others. As scientists have discovered more theropods closely related to birds, the previously clear distinction between non-birds and birds has become blurred. Recent discoveries in the
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
Province of northeast China, which demonstrate many small theropod
feathered dinosaur A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been su ...
s, contribute to this ambiguity. The consensus view in contemporary
palaeontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
is that the flying theropods, or
avialans Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally use ...
, are the closest relatives of the
deinonychosaur Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica,Case, J.A., Mar ...
s, which include dromaeosaurids and
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
s. Together, these form a group called
Paraves Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avial ...
. Some basal members of Deinonychosauria, such as ''
Microraptor ''Microraptor'' (Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. They dat ...
'', have features which may have enabled them to glide or fly. The most basal deinonychosaurs were very small. This evidence raises the possibility that the ancestor of all paravians may have been
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
, have been able to glide, or both. Unlike ''Archaeopteryx'' and the non-avialan feathered dinosaurs, who primarily ate meat, recent studies suggest that the first avialans were
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s. The
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
''Archaeopteryx'' is well known as one of the first
transitional fossil A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
s to be found, and it provided support for the
theory of evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
in the late 19th century. ''Archaeopteryx'' was the first fossil to display both clearly traditional reptilian characteristics—teeth, clawed fingers, and a long, lizard-like tail—as well as wings with flight feathers similar to those of modern birds. It is not considered a direct ancestor of birds, though it is possibly closely related to the true ancestor.


Early evolution

Over 40% of key traits found in modern birds evolved during the 60 million year transition from the earliest bird-line archosaurs to the first maniraptoromorphs, i.e. the first dinosaurs closer to living birds than to ''
Tyrannosaurus rex ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
''. The loss of osteoderms otherwise common in archosaurs and acquisition of primitive feathers might have occurred early during this phase. After the appearance of Maniraptoromorpha, the next 40 million years marked a continuous reduction of body size and the accumulation of
neotenic Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
(juvenile-like) characteristics.
Hypercarnivory A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant examp ...
became increasingly less common while braincases enlarged and forelimbs became longer. The
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind. Etymology The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
evolved into complex,
pennaceous feather The pennaceous feather is a type of feather present in most modern birds and in some other species of maniraptoriform dinosaurs. Description A pennaceous feather has a stalk or quill. Its basal part, called a ''calamus'', is embedded in the skin ...
s. The oldest known paravian (and probably the earliest avialan) fossils come from the
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian- Oxfordian stages). It is known for its exceptionally preserved fossils, including those of ...
of China, which has been dated to the late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period ( Oxfordian stage), about 160 million years ago. The avialan species from this time period include ''
Anchiornis huxleyi ''Anchiornis'' is a genus of small, four-winged paravian dinosaurs, with only one known species, the type species ''Anchiornis huxleyi'', named for its similarity to modern birds. The Latin name ''Anchiornis'' derives from a Greek word meaning " ...
'', '' Xiaotingia zhengi'', and ''
Aurornis xui ''Aurornis'' is an extinct genus of anchiornithid theropod dinosaurs from the Jurassic period of China. The genus ''Aurornis'' contains a single known species, ''Aurornis xui'' (). ''Aurornis xui'' may be the most basal ("primitive") avialan d ...
''. The well-known probable early avialan, ''Archaeopteryx'', dates from slightly later Jurassic rocks (about 155 million years old) from
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. Many of these early avialans shared unusual anatomical features that may be ancestral to modern birds but were later lost during bird evolution. These features include enlarged claws on the second toe which may have been held clear of the ground in life, and long feathers or "hind wings" covering the hind limbs and feet, which may have been used in aerial manoeuvreing. Avialans diversified into a wide variety of forms during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period. Many groups retained primitive characteristics, such as clawed wings and teeth, though the latter were lost independently in a number of avialan groups, including modern birds (Aves). Increasingly stiff tails (especially the outermost half) can be seen in the evolution of maniraptoromorphs, and this process culminated in the appearance of the
pygostyle Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compon ...
, an ossification of fused tail vertebrae. In the late Cretaceous, about 100 million years ago, the ancestors of all modern birds evolved a more open pelvis, allowing them to lay larger eggs compared to body size. Around 95 million years ago, they evolved a better sense of smell. A third stage of bird evolution starting with
Ornithothoraces Ornithothoraces is a group of avialans that includes all enantiornithes ("opposite birds") and the euornithes ("true birds"), which includes modern birds and their closest ancestors. The name Ornithothoraces means "bird thoraxes". This refers to ...
(the "bird-chested" avialans) can be associated with the refining of aerodynamics and flight capabilities, and the loss or co-ossification of several skeletal features. Particularly significant are the development of an enlarged, keeled sternum and the
alula The alula , or bastard wing, (plural ''alulae'') is a small projection on the anterior edge of the wing of modern birds and a few non-avian dinosaurs. The word is Latin and means "winglet"; it is the diminutive of ''ala'', meaning "wing". The al ...
, and the loss of grasping hands.


Early diversity of bird ancestors

The first large, diverse lineage of short-tailed avialans to evolve were the
Enantiornithes The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and cla ...
, or "opposite birds", so named because the construction of their shoulder bones was in reverse to that of modern birds. Enantiornithes occupied a wide array of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s, from sand-probing shorebirds and fish-eaters to tree-dwelling forms and seed-eaters. While they were the dominant group of avialans during the Cretaceous period, enantiornithes became extinct along with many other dinosaur groups at the end of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
era. Many species of the second major avialan lineage to diversify, the
Euornithes Euornithes (from Greek ' meaning "true birds") is a natural group which includes the most recent common ancestor of all avialans closer to modern birds than to ''Sinornis''. Description Clarke ''et al''. (2006) found that the most primitive know ...
(meaning "true birds", because they include the ancestors of modern birds), were semi-aquatic and specialised in eating fish and other small aquatic organisms. Unlike the Enantiornithes, which dominated land-based and arboreal habitats, most early euornithes lacked perching adaptations and seem to have included shorebird-like species, waders, and swimming and diving species. The latter included the superficially
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century, m ...
-like ''
Ichthyornis ''Ichthyornis'' (meaning "fish bird", after its fish-like vertebrae) is an extinct genus of toothy seabird-like ornithuran from the late Cretaceous period of North America. Its fossil remains are known from the chalks of Alberta, Alabama, Kansa ...
'' and the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic. The early euornithes also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithes retained teeth in other parts of the jaws). Euornithes also included the first avialans to develop true
pygostyle Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compon ...
and a fully mobile fan of tail feathers, which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight. A study on
mosaic evolution Mosaic evolution (or modular evolution) is the concept, mainly from palaeontology, that evolutionary change takes place in some body parts or systems without simultaneous changes in other parts. Another definition is the "evolution of characters ...
in the avian skull found that the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of all Neornithes might have had a beak similar to that of the modern
hook-billed vanga The hook-billed vanga (''Vanga curvirostris'') is a species of bird in the family Vangidae. It is endemic to Madagascar. Its natural habitats are subtropical or tropical dry forest, subtropical or tropical moist lowland forest, and subtropical o ...
and a skull similar to that of the
Eurasian golden oriole The Eurasian golden oriole (''Oriolus oriolus'') also called the common golden oriole, is the only member of the Old World oriole family of passerine birds breeding in Northern Hemisphere temperate regions. It is a summer migrant in Europe and ...
. As both species are small aerial and canopy foraging omnivores, a similar ecological niche was inferred for this hypothetical ancestor.


Diversification of modern birds

All modern birds lie within the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
Aves (alternately Neornithes), which has two subdivisions: the
Palaeognathae Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae contai ...
, which includes the flightless
ratite A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites. The systematics ...
s (such as the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
es) and the weak-flying
tinamou Tinamous () form an order of birds called Tinamiformes (), comprising a single family called Tinamidae (), divided into two distinct subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" come ...
s, and the extremely diverse
Neognathae Neognathae (; ) is a infraclass of birds, called neognaths, within the class Aves of the clade Archosauria. Neognathae includes the majority of living birds; the exceptions being the tinamous and the flightless ratites, which belong instead to ...
, containing all other birds. These two subdivisions have variously been given the
rank Rank is the relative position, value, worth, complexity, power, importance, authority, level, etc. of a person or object within a ranking, such as: Level or position in a hierarchical organization * Academic rank * Diplomatic rank * Hierarchy * H ...
of
superorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
, cohort, or infraclass. Depending on the
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
viewpoint, the number of known living bird species varies anywhere from 9,800 to 10,758. The discovery of ''
Vegavis ''Vegavis'' is a genus of extinct bird that lived during the Late Cretaceous (Maastrichtian stage) of Antarctica, some 68 to 66 mya. Among modern birds, most studies show that ''Vegavis'' is most closely related to ducks and geese (Anatidae), b ...
'' from the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
, the last stage of the Late Cretaceous proved that the diversification of modern birds started before the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
era. The affinities of an earlier fossil, the possible
galliform Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are often ...
''
Austinornis ''Austinornis'' is a genus of prehistoric birds related to Galliformes. It is known from a fossil partial tarsometatarsus from the Late Cretaceous of Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central regio ...
lentus'', dated to about 85 million years ago, are still too controversial to provide a fossil evidence of modern bird diversification. In 2020, ''
Asteriornis ''Asteriornis'' ("Asteria (Titaness), Asteria's bird") is an extinct genus of bird from the Late Cretaceous of Belgium which is known from a single species, ''Asteriornis maastrichtensis''. It was closely related to birds of the extant superorder ...
'' from the Maastrichtian was described, it appears to be a close relative of Galloanserae, the earliest diverging lineage within Neognathae. Most studies agree on a Cretaceous age for the most recent common ancestor of modern birds but estimates range from the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
to the latest
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
. Similarly, there is no agreement on whether most of the early diversification of modern birds occurred before or after the Cretaceous–Palaeogene extinction event. This disagreement is in part caused by a divergence in the evidence; most molecular dating studies suggests a Cretaceous
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid ...
, while fossil evidence points to a Cenozoic radiation (the so-called 'rocks' versus 'clocks' controversy). Previous attempts to reconcile molecular and fossil evidence have proved controversial, but more recent estimates, using a more comprehensive sample of fossils and a new way of calibrating
molecular clocks The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotid ...
, showed that while according to some studies, modern birds originated early in the Late Cretaceous in Western
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
, a pulse of diversification in all major groups occurred around the Cretaceous–Palaeogene extinction event. Modern birds expanded from West Gondwana to the Laurasia through two routes. One route was an Antarctic interchange in the Paleogene. This can be confirmed with the presence of multiple avian groups in Australia and New Zealand. The other route was probably through North America, via land bridges, during the Paleocene. This allowed the expansion and diversification of Neornithes into the Holarctic and Paleotropics. On the other hand, the occurrence of ''Asteriornis'' in the Northern Hemisphere challenges biogeographical hypotheses of a Gondwanan origin of crown birds.


Classification of bird orders

Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
of modern bird relationships based on Braun & Kimball (2021) The classification of birds is a contentious issue. Sibley and Ahlquist's ''Phylogeny and Classification of Birds'' (1990) is a landmark work on the subject. Most evidence seems to suggest the assignment of orders is accurate, but scientists disagree about the relationships among the orders themselves; evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem, but no strong consensus has emerged. More recently, new fossil and molecular evidence is providing an increasingly clear picture of the evolution of modern bird orders.


Genomics

, the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
had been sequenced for only two birds, the
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adult m ...
and the
zebra finch The zebra finches are two species of estrildid finch in the genus ''Taeniopygia'' found in Australia and Indonesia. They are seed-eaters that travel in large flocks. The species are: Previously, both species were classified as a single species ...
. the genomes of 542 species of birds had been completed. At least one genome has been sequenced from every order. These include at least one species in about 90% of extant avian families (218 out of 236 families recognised by the ''Howard and Moore Checklist''). Being able to sequence and compare whole genomes gives researchers many types of information, about genes, the DNA that regulates the genes, and their evolutionary history. This has led to reconsideration of some of the classifications that were based solely on the identification of protein-coding genes. Waterbirds such as
pelicans Pelicans (genus ''Pelecanus'') are a genus of large water birds that make up the family Pelecanidae. They are characterized by a long beak and a large throat pouch used for catching prey and draining water from the scooped-up contents before s ...
and
flamingos Flamingos or flamingoes are a type of wading bird in the family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas (including the Caribbean) ...
, for example, may have in common specific adaptations suited to their environment that were developed independently.


Distribution

Birds live and breed in most terrestrial habitats and on all seven continents, reaching their southern extreme in the
snow petrel The snow petrel (''Pagodroma nivea'') is the only member of the genus ''Pagodroma.'' It is one of only three birds that has been seen at the Geographic South Pole, along with the Antarctic petrel and the south polar skua, which have the most so ...
's breeding colonies up to inland in
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
. The highest bird
diversity Diversity, diversify, or diverse may refer to: Business *Diversity (business), the inclusion of people of different identities (ethnicity, gender, age) in the workforce *Diversity marketing, marketing communication targeting diverse customers * ...
occurs in tropical regions. It was earlier thought that this high diversity was the result of higher
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
rates in the tropics; however recent studies found higher speciation rates in the high latitudes that were offset by greater
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
rates than in the tropics. Many species migrate annually over great distances and across oceans; several families of birds have adapted to life both on the world's oceans and in them, and some
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
species come ashore only to breed, while some
penguin Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s have been recorded diving up to deep. Many bird species have established breeding populations in areas to which they have been introduced by humans. Some of these introductions have been deliberate; the
ring-necked pheasant The common pheasant (''Phasianus colchicus'') is a bird in the pheasant family (Phasianidae). The genus name comes from Latin ''phasianus'', "pheasant". The species name ''colchicus'' is Latin for "of Colchis" (modern day Georgia), a country on ...
, for example, has been introduced around the world as a
game bird Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are ofte ...
. Others have been accidental, such as the establishment of wild
monk parakeet The monk parakeet (''Myiopsitta monachus''), also known as the Quaker parrot, is a species of true parrot in the family Psittacidae. It is a small, bright-green parrot with a greyish breast and greenish-yellow abdomen. Its average lifespan is 20 ...
s in several North American cities after their escape from captivity. Some species, including
cattle egret The cattle egret (''Bubulcus ibis'') is a cosmopolitan species of heron (family Ardeidae) found in the tropics, subtropics, and warm-temperate zones. It is the only member of the monotypic genus ''Bubulcus'', although some authorities regard it ...
,
yellow-headed caracara The yellow-headed caracara (''Daptrius chimachima'') is a bird of prey in the family Falconidae. It is found in tropical and subtropical South America and the southern portion of Central America. Unlike the falcons in the same family, the caraca ...
and
galah The galah (; ''Eolophus roseicapilla''), also known as the pink and grey cockatoo or rose-breasted cockatoo, is the only species within genus ''Eolophus'' of the cockatoo family. Found throughout Australia, it is among the most common of the co ...
, have spread naturally far beyond their original ranges as
agricultural expansion Agricultural expansion describes the growth of agricultural land (arable land, pastures, etc.) especially in the 20th and 21st centuries. The agricultural expansion is often explained as a direct consequence of the global increase in food and ene ...
created alternative habitats although modern practices of intensive agriculture have negatively impacted farmland bird populations.


Anatomy and physiology

Compared with other vertebrates, birds have a
body plan A body plan, ( ), or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually applied to animals, envisages a "blueprin ...
that shows many unusual adaptations, mostly to facilitate
flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...
.


Skeletal system

The skeleton consists of very lightweight bones. They have large air-filled cavities (called pneumatic cavities) which connect with the
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...
. The skull bones in adults are fused and do not show
cranial sutures In anatomy, fibrous joints are joints connected by fibrous tissue, consisting mainly of collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of c ...
. The orbital cavities that house the eyeballs are large and separated from each other by a bony
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatri ...
(partition). The spine has cervical, thoracic, lumbar and caudal regions with the number of cervical (neck) vertebrae highly variable and especially flexible, but movement is reduced in the anterior
thoracic vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebra (anatomy), vertebrae and they are intermediate in size b ...
and absent in the later vertebrae. The last few are fused with the
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
to form the synsacrum. The ribs are flattened and the
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sh ...
is keeled for the attachment of flight muscles except in the flightless bird orders. The forelimbs are modified into wings. The wings are more or less developed depending on the species; the only known groups that lost their wings are the
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
and
elephant bird Elephant birds are members of the extinct ratite family Aepyornithidae, made up of flightless birds that once lived on the island of Madagascar. They are thought to have become extinct around 1000-1200 CE, probably as a result of human activity. ...
s.


Excretory system

Like the
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, birds are primarily
uricotelic Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, w ...
, that is, their
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
s extract
nitrogenous waste Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, ...
from their bloodstream and excrete it as
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown of ...
, instead of
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important r ...
or
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
, through the ureters into the intestine. Birds do not have a
urinary bladder The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...
or external urethral opening and (with exception of the
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There are ...
) uric acid is excreted along with faeces as a semisolid waste. However, birds such as hummingbirds can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. They also excrete
creatine Creatine ( or ) is an organic compound with the nominal formula (H2N)(HN)CN(CH3)CH2CO2H. It exists in various modifications (tautomers) in solution. Creatine is found in vertebrates where it facilitates recycling of adenosine triphosphate ( ...
, rather than
creatinine Creatinine (; ) is a breakdown product of creatine phosphate from muscle and protein metabolism. It is released at a constant rate by the body (depending on muscle mass). Biological relevance Serum creatinine (a blood measurement) is an import ...
like mammals. This material, as well as the output of the intestines, emerges from the bird's
cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...
. The cloaca is a multi-purpose opening: waste is expelled through it, most birds mate by joining cloaca, and females lay eggs from it. In addition, many species of birds regurgitate pellets. It is a common but not universal feature of
altricial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
nestlings (born helpless, under constant parental care) that instead of excreting directly into the nest, they produce a
fecal sac A fecal sac (also spelled faecal sac) is a mucous membrane, generally white or clear with a dark end, that surrounds the feces of some species of nestling birds. It allows parent birds to more easily remove fecal material from the nest. The nes ...
. This is a mucus-covered pouch that allows parents to either dispose of the waste outside the nest or to recycle the waste through their own digestive system.


Reproductive system

Males within
Palaeognathae Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae contai ...
(with the exception of the
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...
s), the
Anseriformes Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which in ...
(with the exception of
screamer The screamers are three South American bird species placed in Family (biology), family Anhimidae. They were thought to be related to the Galliformes because of similar beak, bills, but are more closely related to ducks (family Anatidae),Todd, F. ...
s), and in rudimentary forms in
Galliformes Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are often ...
(but fully developed in
Cracidae The chachalacas, guans and curassows are birds in the family Cracidae. These are species of tropical and subtropical Central and South America. The range of one species, the plain chachalaca, just reaches southernmost parts of Texas in the Unite ...
) possess a
penis A penis (plural ''penises'' or ''penes'' () is the primary sexual organ that male animals use to inseminate females (or hermaphrodites) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males do n ...
, which is never present in
Neoaves Neoaves is a clade that consists of all modern birds (Neornithes or Aves) with the exception of Paleognathae (ratites and kin) and Galloanserae (ducks, chickens and kin). Almost 95% of the roughly 10,000 known species of extant birds belong to ...
. The length is thought to be related to
sperm competition Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and ...
. For male birds to get an erection, they depends on lymphatic fluid instead of blood. When not copulating, it is hidden within the
proctodeum A proctodeum is the back ectodermal part of an alimentary canal. It is created during embryogenesis by a folding of the outer body wall. It will form the lower part of the anal canal The anal canal is the part that connects the rectum to the ...
compartment within the cloaca, just inside the vent. Female birds have sperm storage tubules that allow sperm to remain viable long after copulation, a hundred days in some species. Sperm from multiple males may
compete Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
through this mechanism. Most female birds have a single
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
and a single
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, o ...
, both on the left side, but there are exceptions: species in at least 16 different orders of birds have two ovaries. Even these species, however, tend to have a single oviduct. It has been speculated that this might be an adaptation to flight, but males have two testes, and it is also observed that the gonads in both sexes decrease dramatically in size outside the breeding season. Also terrestrial birds generally have a single ovary, as does the
platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal Endemic (ecology), endemic to Eastern states of Australia, eastern Australia, including Tasmania. The platypu ...
, an egg-laying mammal. A more likely explanation is that the egg develops a shell while passing through the oviduct over a period of about a day, so that if two eggs were to develop at the same time, there would be a risk to survival. While rare, mostly abortive,
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
is not unknown in birds and eggs can be
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, automictic and results in male offspring. Birds are solely
gonochoric In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric. Gonochorism c ...
. Meaning they have two sexes: either
female Female (Venus symbol, symbol: ♀) is the sex of an organism that produces the large non-motile ovum, ova (egg cells), the type of gamete (sex cell) that fuses with the Sperm, male gamete during sexual reproduction. A female has larger gamet ...
or
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
. The sex of birds is determined by the Z and W sex chromosomes, rather than by the X and Y chromosomes present in
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. Male birds have two Z chromosomes (ZZ), and female birds have a W chromosome and a Z chromosome (WZ). In nearly all species of birds, an individual's sex is determined at fertilisation. However, one 2007 study claimed to demonstrate
temperature-dependent sex determination Temperature-dependent sex determination (TSD) is a type of environmental sex determination in which the temperatures experienced during embryonic/larval development determine the sex of the offspring. It is only observed in reptiles and teleost fish ...
among the
Australian brushturkey The Australian brushturkey or Australian brush-turkey or gweela (''Alectura lathami''), also frequently called the scrub turkey or bush turkey, is a common, widespread species of mound-building bird from the family Megapodiidae found in eastern ...
, for which higher temperatures during incubation resulted in a higher female-to-male
sex ratio The sex ratio (or gender ratio) is usually defined as the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. Many species devia ...
. This, however, was later proven to not be the case. These birds do not exhibit temperature-dependent sex determination, but temperature-dependent sex mortality.


Respiratory and circulatory systems

Birds have one of the most complex
respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...
s of all animal groups. Upon inhalation, 75% of the fresh air bypasses the lungs and flows directly into a posterior
air sac Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence ...
which extends from the lungs and connects with air spaces in the bones and fills them with air. The other 25% of the air goes directly into the lungs. When the bird exhales, the used air flows out of the lungs and the stored fresh air from the posterior air sac is simultaneously forced into the lungs. Thus, a bird's lungs receive a constant supply of fresh air during both inhalation and exhalation. Sound production is achieved using the
syrinx In classical Greek mythology, Syrinx (Greek Σύριγξ) was a nymph and a follower of Artemis, known for her chastity. Pursued by the amorous god Pan, she ran to a river's edge and asked for assistance from the river nymphs. In answer, sh ...
, a muscular chamber incorporating multiple tympanic membranes which diverges from the lower end of the trachea; the trachea being elongated in some species, increasing the volume of vocalisations and the perception of the bird's size. In birds, the main arteries taking blood away from the heart originate from the right
aortic arch The aortic arch, arch of the aorta, or transverse aortic arch () is the part of the aorta between the ascending and descending aorta. The arch travels backward, so that it ultimately runs to the left of the trachea. Structure The aorta begins a ...
(or pharyngeal arch), unlike in the mammals where the left aortic arch forms this part of the
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes ...
. The postcava receives blood from the limbs via the renal portal system. Unlike in mammals, the circulating
red blood cells Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek language, Greek ''erythros'' for "red" and ''k ...
in birds retain their
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...
.


Heart type and features

The avian circulatory system is driven by a four-chambered, myogenic heart contained in a fibrous pericardial sac. This pericardial sac is filled with a
serous fluid In physiology, serous fluid or serosal fluid (originating from the Medieval Latin word ''serosus'', from Latin ''serum'') is any of various body fluids resembling Serum (blood), serum, that are typically pale yellow or transparent and of a benign ...
for lubrication. The heart itself is divided into a right and left half, each with an
atrium Atrium may refer to: Anatomy * Atrium (heart), an anatomical structure of the heart * Atrium, the genital structure next to the genital aperture in the reproductive system of gastropods * Atrium of the ventricular system of the brain * Pulmona ...
and ventricle. The atrium and ventricles of each side are separated by
atrioventricular valves A heart valve is a one-way valve that allows blood to flow in one direction through the chambers of the heart. Four valves are usually present in a mammalian heart and together they determine the pathway of blood flow through the heart. A heart v ...
which prevent back flow from one chamber to the next during contraction. Being myogenic, the heart's pace is maintained by pacemaker cells found in the sinoatrial node, located on the right atrium. The
sinoatrial node The sinoatrial node (also known as the sinuatrial node, SA node or sinus node) is an oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of cells known as pacemaker cells. The sinus node is approximat ...
uses calcium to cause a depolarising
signal transduction pathway Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellular ...
from the atrium through right and left atrioventricular bundle which communicates contraction to the ventricles. The avian heart also consists of muscular arches that are made up of thick bundles of muscular layers. Much like a mammalian heart, the avian heart is composed of
endocardial The endocardium is the innermost layer of tissue that lines the chambers of the heart. Its cells are embryologically and biologically similar to the endothelial cells that line blood vessels. The endocardium also provides protection to the va ...
,
myocardial Cardiac muscle (also called heart muscle, myocardium, cardiomyocytes and cardiac myocytes) is one of three types of vertebrate muscle tissues, with the other two being skeletal muscle and smooth muscle. It is an involuntary, striated muscle that ...
and epicardial layers. The atrium walls tend to be thinner than the ventricle walls, due to the intense ventricular contraction used to pump oxygenated blood throughout the body. Avian hearts are generally larger than mammalian hearts when compared to body mass. This adaptation allows more blood to be pumped to meet the high metabolic need associated with flight.


Organisation

Birds have a very efficient system for diffusing oxygen into the blood; birds have a ten times greater surface area to
gas exchange Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
volume than mammals. As a result, birds have more blood in their capillaries per unit of volume of lung than a mammal. The arteries are composed of thick elastic muscles to withstand the pressure of the ventricular contractions, and become more rigid as they move away from the heart. Blood moves through the arteries, which undergo
vasoconstriction Vasoconstriction is the narrowing of the blood vessels resulting from contraction of the muscular wall of the vessels, in particular the large arteries and small arterioles. The process is the opposite of vasodilation, the widening of blood vessel ...
, and into arterioles which act as a transportation system to distribute primarily oxygen as well as nutrients to all tissues of the body. As the arterioles move away from the heart and into individual organs and tissues they are further divided to increase surface area and slow blood flow. Blood travels through the arterioles and moves into the capillaries where gas exchange can occur. Capillaries are organised into capillary beds in tissues; it is here that blood exchanges oxygen for carbon dioxide waste. In the capillary beds, blood flow is slowed to allow maximum
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
of oxygen into the tissues. Once the blood has become deoxygenated, it travels through venules then veins and back to the heart. Veins, unlike arteries, are thin and rigid as they do not need to withstand extreme pressure. As blood travels through the venules to the veins a funneling occurs called
vasodilation Vasodilation is the widening of blood vessels. It results from relaxation of smooth muscle cells within the vessel walls, in particular in the large veins, large arteries, and smaller arterioles. The process is the opposite of vasoconstriction, ...
bringing blood back to the heart. Once the blood reaches the heart, it moves first into the right atrium, then the right ventricle to be pumped through the lungs for further gas exchange of carbon dioxide waste for oxygen. Oxygenated blood then flows from the lungs through the left atrium to the left ventricle where it is pumped out to the body.


Nervous system

The
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
is large relative to the bird's size. The most developed part of the brain is the one that controls the flight-related functions, while the
cerebellum The cerebellum (Latin for "little brain") is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebel ...
coordinates movement and the
cerebrum The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. In ...
controls behaviour patterns, navigation, mating and
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materia ...
building. Most birds have a poor
sense of smell The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
with notable exceptions including
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...
s,
New World vulture The New World vulture or condor family, Cathartidae, contains seven extant species in five genera. It includes five extant vultures and two extant condors found in warm and temperate areas of the Americas. The "New World" vultures were widespread ...
s and tubenoses. The avian
visual system The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the a ...
is usually highly developed. Water birds have special flexible lenses, allowing accommodation for vision in air and water. Some species also have dual
fovea Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure. Human anatomy *Fovea centralis of the retina * Fovea buccalis or Dimple * Fovea of the femoral head * Trochlear fovea of the fr ...
. Birds are tetrachromatic, possessing
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
(UV) sensitive
cone cell Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...
s in the eye as well as green, red and blue ones. They also have double cones, likely to mediate achromatic vision. Many birds show plumage patterns in
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
that are invisible to the human eye; some birds whose sexes appear similar to the naked eye are distinguished by the presence of ultraviolet reflective patches on their feathers. Male
blue tit The Eurasian blue tit (''Cyanistes caeruleus'') is a small passerine bird in the tit family, Paridae. It is easily recognisable by its blue and yellow plumage and small size. Eurasian blue tits, usually resident and non-migratory birds, are ...
s have an ultraviolet reflective crown patch which is displayed in courtship by posturing and raising of their nape feathers. Ultraviolet light is also used in foraging—
kestrel The term kestrel (from french: crécerelle, derivative from , i.e. ratchet) is the common name given to several species of predatory birds from the falcon genus ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviou ...
s have been shown to search for prey by detecting the UV reflective urine trail marks left on the ground by rodents. With the exception of pigeons and a few other species, the eyelids of birds are not used in blinking. Instead the eye is lubricated by the
nictitating membrane The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
, a third eyelid that moves horizontally. The nictitating membrane also covers the eye and acts as a
contact lens Contact lenses, or simply contacts, are thin lenses placed directly on the surface of the eyes. Contact lenses are ocular prosthetic devices used by over 150 million people worldwide, and they can be worn to correct vision or for cosmetic ...
in many aquatic birds. The bird
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
has a fan shaped blood supply system called the pecten. Eyes of most birds are large, not very round and capable of only limited movement in the orbits, typically 10–20°. Birds with eyes on the sides of their heads have a wide
visual field The visual field is the "spatial array of visual sensations available to observation in introspectionist psychological experiments". Or simply, visual field can be defined as the entire area that can be seen when an eye is fixed straight at a point ...
, while birds with eyes on the front of their heads, such as owls, have
binocular vision In biology, binocular vision is a type of vision in which an animal has two eyes capable of facing the same direction to perceive a single three-dimensional image of its surroundings. Binocular vision does not typically refer to vision where an ...
and can estimate the
depth of field The depth of field (DOF) is the distance between the nearest and the furthest objects that are in acceptably sharp focus in an image captured with a camera. Factors affecting depth of field For cameras that can only focus on one object dist ...
. The avian
ear An ear is the organ that enables hearing and, in mammals, body balance using the vestibular system. In mammals, the ear is usually described as having three parts—the outer ear, the middle ear and the inner ear. The outer ear consists of ...
lacks external
pinnae The auricle or auricula is the visible part of the ear that is outside the head. It is also called the pinna (Latin for "wing" or " fin", plural pinnae), a term that is used more in zoology. Structure The diagram shows the shape and location ...
but is covered by feathers, although in some birds, such as the ''
Asio ''Asio'' is a genus of typical owls, or true owls, in the family Strigidae. This group has representatives over most of the planet, and the short-eared owl is one of the most widespread of all bird species, breeding in Europe, Asia, North and S ...
'', ''
Bubo A bubo (Greek βουβών, ''boubṓn'', 'groin') is adenitis or inflammation of the lymph nodes and is an example of reactive lymphadenopathy. Classification Buboes are a symptom of bubonic plague and occur as painful swellings in the thigh ...
'' and ''
Otus Otus may refer to: * Otus (education), a K-12 educational technology company * HMS ''Otus'', two ships in the Royal Navy * ''Otus'' (bird), genus of owls * Otus (mythology), giant in Greek mythology, brother of Ephialtes, one of Aloadae * Otus ...
''
owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...
s, these feathers form tufts which resemble ears. The
inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
has a
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus. A core component of the cochlea is the Organ of Corti, the sensory org ...
, but it is not spiral as in mammals.


Defence and intraspecific combat

A few species are able to use chemical defences against predators; some
Procellariiformes Procellariiformes is an order of seabirds that comprises four families: the albatrosses, the petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still called tubenoses in English, procellariiforms are of ...
can eject an unpleasant
stomach oil Stomach oil is the light oil composed of neutral dietary lipids found in the proventriculus (fore-gut) of birds in the order Procellariiformes. All albatrosses, procellarids (gadfly petrels and shearwaters) and northern and austral storm petrel ...
against an aggressor, and some species of
pitohui The pitohuis are bird species endemic to New Guinea. The onomatopoeic name is thought to be derived from that used by New Guineans from nearby Dorey (Manokwari), but it is also used as the name of a genus '' Pitohui'' which was established by the ...
s from
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
have a powerful
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature ner ...
in their skin and feathers. A lack of field observations limit our knowledge, but intraspecific conflicts are known to sometimes result in injury or death. The screamers ( Anhimidae), some jacanas ('' Jacana'', '' Hydrophasianus''), the spur-winged goose ('' Plectropterus''), the torrent duck (''
Merganetta The torrent duck (''Merganetta armata'') is a member of the duck, goose and swan family Anatidae. It is the only member of the genus ''Merganetta''. It is placed in the shelduck subfamily Tadorninae after the "perching duck" assemblage to which i ...
'') and nine species of lapwing (''
Vanellus ''Vanellus'' is the genus of waders which provisionally contains all lapwings except red-kneed dotterel, ''Erythrogonys cinctus''. The name "''vanellus''" is Latin for "little fan", ''vanellus'' being the diminutive of ''vannus'' ("winnowing fan ...
'') use a sharp spur on the wing as a weapon. The steamer ducks (''
Tachyeres The steamer ducks are a genus (''Tachyeres'') of ducks in the family Anatidae. All of the four species occur at the southern cone of South America in Chile and Argentina, and all except the flying steamer duck are flightless; even this one spec ...
''), geese and swans (''
Anserinae The Anserinae are a subfamily in the waterfowl family Anatidae. It includes the swans and true geese. Under alternative systematical concepts (see e.g., Terres & NAS, 1991), it is split into two subfamilies, the Anserinae contain the geese and ...
''), the solitaire ('' Pezophaps''), sheathbills (''
Chionis The sheathbills are a family of birds, Chionidae. Classified in the wader order Charadriiformes, the family consists of one genus, ''Chionis'' with two species. They breed on subantarctic islands and the Antarctic Peninsula, and the snowy sheat ...
''), some guans ('' Crax'') and stone curlews (''
Burhinus ''Burhinus'' is a genus of birds in the family Burhinidae. This family also contains the genus ''Esacus''.del Hoyo J, Elliott A, Sargatal J (1996) ''Handbook of the Birds of the World'', ''vol 3.'' Lynx, Barcelona The genus name ''Burhinus'' com ...
'') use a bony knob on the
alula The alula , or bastard wing, (plural ''alulae'') is a small projection on the anterior edge of the wing of modern birds and a few non-avian dinosaurs. The word is Latin and means "winglet"; it is the diminutive of ''ala'', meaning "wing". The al ...
r metacarpal to punch and hammer opponents. The jacanas ''
Actophilornis ''Actophilornis'' is a genus of jacana. It contains two species restricted to Africa and its surrounding islands. Taxonomy The genus ''Actophilornis'' was introduced in 1925 by the American ornithologist Harry C. Oberholser with the African ...
'' and ''
Irediparra The comb-crested jacana (''Irediparra gallinacea''), also known as the lotusbird or lilytrotter, is the only species of jacana in the genus ''Irediparra''. Like other jacana species, it is adapted to the floating vegetation of tropical freshwat ...
'' have an expanded, blade-like radius. The extinct '' Xenicibis'' was unique in having an elongate forelimb and massive hand which likely functioned in combat or defence as a jointed club or flail.
Swans Swans are birds of the family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe Cygnini. Someti ...
, for instance, may strike with the bony spurs and bite when defending eggs or young.


Feathers, plumage, and scales

Feathers are a feature characteristic of birds (though also present in some dinosaurs not currently considered to be true birds). They facilitate
flight Flight or flying is the process by which an object moves through a space without contacting any planetary surface, either within an atmosphere (i.e. air flight or aviation) or through the vacuum of outer space (i.e. spaceflight). This can be a ...
, provide insulation that aids in
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
, and are used in display, camouflage, and signalling. There are several types of feathers, each serving its own set of purposes. Feathers are epidermal growths attached to the skin and arise only in specific tracts of skin called pterylae. The distribution pattern of these feather tracts (pterylosis) is used in taxonomy and systematics. The arrangement and appearance of feathers on the body, called
plumage Plumage ( "feather") is a layer of feathers that covers a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between species and subspecies and may vary with age classes. Within species, ...
, may vary within species by age,
social status Social status is the level of social value a person is considered to possess. More specifically, it refers to the relative level of respect, honour, assumed competence, and deference accorded to people, groups, and organizations in a society. Stat ...
, and
sex Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones ( ova, of ...
. Plumage is regularly
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
ed; the standard plumage of a bird that has moulted after breeding is known as the "" plumage, or—in the
Humphrey–Parkes terminology Humphrey–Parkes terminology is a system of nomenclature for the plumage of birds. Before the Humphrey–Parkes system, plumage was named after the belief that a certain plumage was breeding plumage and others were not. However, as this system did ...
—"basic" plumage; breeding plumages or variations of the basic plumage are known under the Humphrey–Parkes system as "" plumages. Moulting is annual in most species, although
some Some may refer to: *''some'', an English word used as a determiner and pronoun; see use of ''some'' *The term associated with the existential quantifier *"Some", a song by Built to Spill from their 1994 album ''There's Nothing Wrong with Love'' *S ...
may have two moults a year, and large birds of prey may moult only once every few years. Moulting patterns vary across species. In passerines,
flight feather Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
s are replaced one at a time with the innermost being the first. When the fifth of sixth primary is replaced, the outermost begin to drop. After the innermost tertiaries are moulted, the starting from the innermost begin to drop and this proceeds to the outer feathers (centrifugal moult). The greater primary are moulted in synchrony with the primary that they overlap. A small number of species, such as ducks and geese, lose all of their flight feathers at once, temporarily becoming flightless. As a general rule, the tail feathers are moulted and replaced starting with the innermost pair. Centripetal moults of tail feathers are however seen in the
Phasianidae The Phasianidae are a family (biology), family of heavy, ground-living birds, which includes pheasants, partridges, junglefowl, chickens, Turkey bird, turkeys, Old World quail, and peafowl. The family includes many of the most popular Game (hunti ...
. The centrifugal moult is modified in the tail feathers of
woodpecker Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...
s and
treecreeper The treecreepers are a family, Certhiidae, of small passerine birds, widespread in wooded regions of the Northern Hemisphere and sub-Saharan Africa. The family contains eleven species in two genera, ''Certhia'' and ''Salpornis''. Their plumage is ...
s, in that it begins with the second innermost pair of feathers and finishes with the central pair of feathers so that the bird maintains a functional climbing tail. The general pattern seen in
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
s is that the primaries are replaced outward, secondaries inward, and the tail from centre outward. Before nesting, the females of most bird species gain a bare brood patch by losing feathers close to the belly. The skin there is well supplied with blood vessels and helps the bird in incubation. Feathers require maintenance and birds preen or groom them daily, spending an average of around 9% of their daily time on this. The bill is used to brush away foreign particles and to apply
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to giv ...
y secretions from the
uropygial gland The uropygial gland, informally known as the preen gland or the oil gland, is a bilobed sebaceous gland possessed by the majority of birds used to distribute the gland's oil through the plumage by means of preening. It is located dorsally at the ...
; these secretions protect the feathers' flexibility and act as an
antimicrobial agent An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
, inhibiting the growth of feather-degrading
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
. This may be supplemented with the secretions of
formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. Es ...
from ants, which birds receive through a behaviour known as
anting Anting () is a town in Jiading District, Shanghai, bordering Kunshan, Jiangsu to the west. It has 96,000 inhabitants and, after the July 2009 merger of Huangdu (), an area of .
, to remove feather parasites. The
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
of birds are composed of the same keratin as beaks, claws, and spurs. They are found mainly on the toes and
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
, but may be found further up on the ankle in some birds. Most bird scales do not overlap significantly, except in the cases of
kingfisher Kingfishers are a family, the Alcedinidae, of small to medium-sized, brightly colored birds in the order Coraciiformes. They have a cosmopolitan distribution, with most species found in the tropical regions of Africa, Asia, and Oceania, ...
s and
woodpecker Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...
s. The scales of birds are thought to be homologous to those of reptiles and mammals.


Flight

Most birds can
fly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...
, which distinguishes them from almost all other vertebrate classes. Flight is the primary means of locomotion for most bird species and is used for searching for food and for escaping from predators. Birds have various adaptations for flight, including a lightweight skeleton, two large flight muscles, the pectoralis (which accounts for 15% of the total mass of the bird) and the supracoracoideus, as well as a modified forelimb (
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...
) that serves as an
aerofoil An airfoil (American English) or aerofoil (British English) is the cross-sectional shape of an object whose motion through a gas is capable of generating significant lift, such as a wing, a sail, or the blades of propeller, rotor, or turbine. ...
. Wing shape and size generally determine a bird's flight style and performance; many birds combine powered, flapping flight with less energy-intensive soaring flight. About 60 extant bird species are
flightless Flightless birds are birds that through evolution lost the ability to fly. There are over 60 extant species, including the well known ratites (ostriches, emu, cassowaries, rheas, and kiwi) and penguins. The smallest flightless bird is the ...
, as were many extinct birds. Flightlessness often arises in birds on isolated islands, most likely due to limited resources and the absence of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
ian land predators. Flightlessness is almost exclusively correlated with
gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
due to an island's inheren condition of isolation. Although flightless, penguins use similar musculature and movements to "fly" through the water, as do some flight-capable birds such as
auk An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...
s,
shearwater Shearwaters are medium-sized long-winged seabirds in the petrel family Procellariidae The family Procellariidae is a group of seabirds that comprises the fulmarine petrels, the gadfly petrels, the diving petrels, the prions, and the shearwa ...
s and
dipper Dippers are members of the genus ''Cinclus'' in the bird family Cinclidae, so-called because of their bobbing or dipping movements. They are unique among passerines for their ability to dive and swim underwater. Taxonomy The genus ''Cinclus'' ...
s.


Behaviour

Most birds are diurnal, but some birds, such as many species of
owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...
s and
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called goatsuckers, due to the ancient folk ta ...
s, are
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
(active during twilight hours), and many coastal
wader 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s feed when the tides are appropriate, by day or night.


Diet and feeding

are varied and often include
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
, fruit, plants, seeds,
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
, and various small animals, including other birds. The
digestive system of birds Bird anatomy, or the physiological structure of birds' bodies, shows many unique adaptations, mostly aiding flight. Birds have a light skeletal system and light but powerful musculature which, along with circulatory and respiratory systems capabl ...
is unique, with a
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydroponic ...
for storage and a
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...
that contains swallowed stones for grinding food to compensate for the lack of teeth. Some species such as pigeons and some psittacine species do not have a
gallbladder In vertebrates, the gallbladder, also known as the cholecyst, is a small hollow organ where bile is stored and concentrated before it is released into the small intestine. In humans, the pear-shaped gallbladder lies beneath the liver, although ...
. Most birds are highly adapted for rapid digestion to aid with flight. Some migratory birds have adapted to use protein stored in many parts of their bodies, including protein from the intestines, as additional energy during migration. (Erratum in ''Proceedings of the Royal Society B'' 267(1461):2567.) Birds that employ many strategies to obtain food or feed on a variety of food items are called generalists, while others that concentrate time and effort on specific food items or have a single strategy to obtain food are considered specialists.
Avian foraging Avian foraging refers to the range of activities and behaviours exhibited by birds in their quest for food. In addition to their unique body adaptations, birds have a range of described behaviours that differ from the foraging behaviours of other a ...
strategies can vary widely by species. Many birds
glean Gleaning is the act of collecting leftover crops from farmers' fields after they have been commercially harvested or on fields where it is not economically profitable to harvest. It is a practice described in the Hebrew Bible that became a legall ...
for insects, invertebrates, fruit, or seeds. Some hunt insects by suddenly attacking from a branch. Those species that seek pest
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s are considered beneficial 'biological control agents' and their presence encouraged in
biological pest control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
programmes. Combined, insectivorous birds eat 400–500 million metric tons of arthropods annually. Nectar feeders such as
hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s,
sunbird Sunbirds and spiderhunters make up the family Nectariniidae of passerine birds. They are small, slender passerines from the Old World, usually with downward-curved bills. Many are brightly coloured, often with iridescent feathers, particularly i ...
s, lories, and lorikeets amongst others have specially adapted brushy tongues and in many cases bills designed to fit
co-adapted In biology, co-adaptation is the process by which two or more species, genes or phenotypic traits undergo adaptation as a pair or group. This occurs when two or more interacting characteristics undergo natural selection together in response to the s ...
flowers.
Kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...
s and
shorebird 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s with long bills probe for invertebrates; shorebirds' varied bill lengths and feeding methods result in the separation of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s.
Loon Loons (North American English) or divers (British / Irish English) are a group of aquatic birds found in much of North America and northern Eurasia. All living species of loons are members of the genus ''Gavia'', family Gaviidae and order Gavi ...
s,
diving duck The diving ducks, commonly called pochards or scaups, are a category of duck which feed by diving beneath the surface of the water. They are part of Anatidae, the diverse and very large family that includes ducks, geese, and swans. The diving d ...
s,
penguin Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s and
auks An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...
pursue their prey underwater, using their wings or feet for propulsion, while aerial predators such as sulids,
kingfisher Kingfishers are a family, the Alcedinidae, of small to medium-sized, brightly colored birds in the order Coraciiformes. They have a cosmopolitan distribution, with most species found in the tropical regions of Africa, Asia, and Oceania, ...
s and
tern Terns are seabirds in the family Laridae that have a worldwide distribution and are normally found near the sea, rivers, or wetlands. Terns are treated as a subgroup of the family Laridae which includes gulls and skimmers and consists of e ...
s plunge dive after their prey.
Flamingo Flamingos or flamingoes are a type of Wader, wading bird in the Family (biology), family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas ...
s, three species of
prion Prions are misfolded proteins that have the ability to transmit their misfolded shape onto normal variants of the same protein. They characterize several fatal and transmissible neurodegenerative diseases in humans and many other animals. It ...
, and some ducks are
filter feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s.
Geese A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...
and
dabbling duck The Anatinae are a subfamily of the family Anatidae ( swans, geese and ducks). Its surviving members are the dabbling ducks, which feed mainly at the surface rather than by diving. The other members of the Anatinae are the extinct moa-nalo, a yo ...
s are primarily grazers. Some species, including
frigatebird Frigatebirds are a family of seabirds called Fregatidae which are found across all tropical and subtropical oceans. The five extant species are classified in a single genus, ''Fregata''. All have predominantly black plumage, long, deeply forked ...
s,
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century, m ...
s, and
skua The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the long-tailed skua, the Arctic skua, and the pomarine skua are called jae ...
s, engage in
kleptoparasitism Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when foo ...
, stealing food items from other birds. Kleptoparasitism is thought to be a supplement to food obtained by hunting, rather than a significant part of any species' diet; a study of
great frigatebird The great frigatebird (''Fregata minor'') is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific (including the Galapagos Islands) and Indian Oceans, as well as a tiny population in the South At ...
s stealing from masked boobies estimated that the frigatebirds stole at most 40% of their food and on average stole only 5%. Other birds are
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s; some of these, like
vulture A vulture is a bird of prey that scavenges on carrion. There are 23 extant species of vulture (including Condors). Old World vultures include 16 living species native to Europe, Africa, and Asia; New World vultures are restricted to North and ...
s, are specialised carrion eaters, while others, like gulls,
corvid Corvidae is a Cosmopolitan distribution, cosmopolitan family (biology), family of oscine passerine birds that contains the crows, ravens, rook (bird), rooks, jackdaws, jays, magpies, treepies, choughs, and nutcracker (bird), nutcrackers. In coll ...
s, or other birds of prey, are opportunists.


Water and drinking

Water is needed by many birds although their mode of excretion and lack of
sweat gland Sweat glands, also known as sudoriferous or sudoriparous glands, , are small tubular structures of the skin that produce sweat. Sweat glands are a type of exocrine gland, which are glands that produce and secrete substances onto an epithelial sur ...
s reduces the physiological demands. Some desert birds can obtain their water needs entirely from moisture in their food. They may also have other adaptations such as allowing their body temperature to rise, saving on moisture loss from evaporative cooling or panting. Seabirds can drink seawater and have
salt gland The salt gland is an organ for excreting excess salts. It is found in the cartilaginous fishes subclass elasmobranchii (sharks, rays, and skates), seabirds, and some reptiles. Salt glands can be found in the rectum of sharks. Birds and reptile ...
s inside the head that eliminate excess salt out of the nostrils. Most birds scoop water in their beaks and raise their head to let water run down the throat. Some species, especially of arid zones, belonging to the
pigeon Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
,
finch The true finches are small to medium-sized passerine birds in the family Fringillidae. Finches have stout conical bills adapted for eating seeds and nuts and often have colourful plumage. They occupy a great range of habitats where they are usua ...
,
mousebird The mousebirds are birds in the order Coliiformes. They are the sister group to the clade Eucavitaves, which includes the Leptosomiformes (the cuckoo roller), Trogoniformes (trogons), Bucerotiformes (hornbills and hoopoes), Piciformes (woodpec ...
,
button-quail Buttonquail or hemipodes are members of a small family of birds, Turnicidae, which resemble, but are unrelated to, the quails of Phasianidae. They inhabit warm grasslands in Asia, Africa, Europe, and Australia. There are 18 species in two genera ...
and
bustard Bustards, including floricans and korhaans, are large, terrestrial birds living mainly in dry grassland areas and on the steppes of the Old World. They range in length from . They make up the family Otididae (, formerly known as Otidae). Bustards ...
families are capable of sucking up water without the need to tilt back their heads. Some desert birds depend on water sources and
sandgrouse Sandgrouse is the common name for Pteroclidae , a family (biology), family of sixteen species of bird, members of the order Pterocliformes . They are traditionally placed in two Genus, genera. The two central Asian species are classified as ''Sy ...
are particularly well known for their daily congregations at waterholes. Nesting sandgrouse and many plovers carry water to their young by wetting their belly feathers. Some birds carry water for chicks at the nest in their crop or regurgitate it along with food. The pigeon family, flamingos and penguins have adaptations to produce a nutritive fluid called
crop milk Crop milk is a secretion from the lining of the crop of parent birds that is regurgitated to young birds. It is found among all pigeons and doves where it is referred to as pigeon milk. An analog to crop milk is also secreted from the esopha ...
that they provide to their chicks.


Feather care

Feathers, being critical to the survival of a bird, require maintenance. Apart from physical wear and tear, feathers face the onslaught of fungi,
ectoparasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
feather mites and
bird lice A bird louse is any chewing louse (small, biting insects) of order Phthiraptera which parasitizes warm-blooded animals, especially birds. Bird lice may feed on feathers, skin, or blood. They have no wings, and their biting mouth parts distingui ...
. The physical condition of feathers are maintained by often with the application of secretions from the . Birds also bathe in water or dust themselves. While some birds dip into shallow water, more aerial species may make aerial dips into water and arboreal species often make use of dew or rain that collect on leaves. Birds of arid regions make use of loose soil to dust-bathe. A behaviour termed as
anting Anting () is a town in Jiading District, Shanghai, bordering Kunshan, Jiangsu to the west. It has 96,000 inhabitants and, after the July 2009 merger of Huangdu (), an area of .
in which the bird encourages ants to run through their plumage is also thought to help them reduce the ectoparasite load in feathers. Many species will spread out their wings and expose them to direct sunlight and this too is thought to help in reducing fungal and ectoparasitic activity that may lead to feather damage.


Migration

Many bird species migrate to take advantage of global differences of
season A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
al temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds,
shorebird 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s, and
waterbird A water bird, alternatively waterbird or aquatic bird, is a bird that lives on or around water. In some definitions, the term ''water bird'' is especially applied to birds in freshwater ecosystems, although others make no distinction from seabi ...
s undertake annual long-distance migrations, usually triggered by the length of daylight as well as weather conditions. These birds are characterised by a breeding season spent in the
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
or
polar region The polar regions, also called the frigid geographical zone, zones or polar zones, of Earth are the regions of the planet that surround its geographical poles (the North Pole, North and South Poles), lying within the polar circles. These high l ...
s and a non-breeding season in the
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs. (Erratum in ''Proceedings of the Royal Society B'' 267(1461):2567.) Migration is highly demanding energetically, particularly as birds need to cross deserts and oceans without refuelling. Landbirds have a flight range of around and shorebirds can fly up to , although the
bar-tailed godwit The bar-tailed godwit (''Limosa lapponica'') is a large and strongly migratory wader in the family Scolopacidae, which feeds on bristle-worms and shellfish on coastal mudflats and estuaries. It has distinctive red breeding plumage, long legs, an ...
is capable of non-stop flights of up to .
Seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s also undertake long migrations, the longest annual migration being those of
sooty shearwater The sooty shearwater (''Ardenna grisea'') is a medium-large shearwater in the seabird family Procellariidae. In New Zealand, it is also known by its Māori name , and as muttonbird, like its relatives the wedge-tailed shearwater (''A. pacificus' ...
s, which nest in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
and
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
and spend the northern summer feeding in the North Pacific off Japan,
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
and
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
, an annual round trip of . Other seabirds disperse after breeding, travelling widely but having no set migration route.
Albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
es nesting in the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the World Ocean, generally taken to be south of 60° S latitude and encircling Antarctica. With a size of , it is regarded as the second-small ...
often undertake circumpolar trips between breeding seasons. Some bird species undertake shorter migrations, travelling only as far as is required to avoid bad weather or obtain food. Irruptive species such as the boreal
finch The true finches are small to medium-sized passerine birds in the family Fringillidae. Finches have stout conical bills adapted for eating seeds and nuts and often have colourful plumage. They occupy a great range of habitats where they are usua ...
es are one such group and can commonly be found at a location in one year and absent the next. This type of migration is normally associated with food availability. Species may also travel shorter distances over part of their range, with individuals from higher latitudes travelling into the existing range of conspecifics; others undertake partial migrations, where only a fraction of the population, usually females and subdominant males, migrates. Partial migration can form a large percentage of the migration behaviour of birds in some regions; in Australia, surveys found that 44% of non-passerine birds and 32% of passerines were partially migratory.
Altitudinal migration Altitudinal migration is a short-distance animal migration from lower altitudes to higher altitudes and back. Altitudinal migrants change their elevation with the seasons making this form of animal migration seasonal. Altitudinal migration can be m ...
is a form of short-distance migration in which birds spend the breeding season at higher altitudes and move to lower ones during suboptimal conditions. It is most often triggered by temperature changes and usually occurs when the normal territories also become inhospitable due to lack of food. Some species may also be nomadic, holding no fixed territory and moving according to weather and food availability.
Parrots Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genus (biology), genera comprising the order (biology), order Psittaciformes (), found mostly in tropics, tropical and subtropics, subtropical regions. The order ...
as a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
are overwhelmingly neither migratory nor sedentary but considered to either be dispersive, irruptive, nomadic or undertake small and irregular migrations. The ability of birds to return to precise locations across vast distances has been known for some time; in an experiment conducted in the 1950s, a
Manx shearwater The Manx shearwater (''Puffinus puffinus'') is a medium-sized shearwater in the seabird family Procellariidae. The scientific name of this species records a name shift: Manx shearwaters were called Manks puffins in the 17th century. Puffin is an ...
released in
Boston Boston (), officially the City of Boston, is the state capital and most populous city of the Commonwealth of Massachusetts, as well as the cultural and financial center of the New England region of the United States. It is the 24th- mo ...
in the United States returned to its colony in
Skomer Skomer () or Skomer Island is an island off the coast of Pembrokeshire, in the community of Marloes and St Brides in west Wales. It is well known for its wildlife: around half the world's population of Manx shearwaters nest on the island, the At ...
, in Wales within 13 days, a distance of . Birds navigate during migration using a variety of methods. For diurnal migrants, the
sun The Sun is the star at the center of the Solar System. It is a nearly perfect ball of hot plasma, heated to incandescence by nuclear fusion reactions in its core. The Sun radiates this energy mainly as light, ultraviolet, and infrared radi ...
is used to navigate by day, and a stellar compass is used at night. Birds that use the sun compensate for the changing position of the sun during the day by the use of an internal clock. Orientation with the stellar compass depends on the position of the
constellation A constellation is an area on the celestial sphere in which a group of visible stars forms Asterism (astronomy), a perceived pattern or outline, typically representing an animal, mythological subject, or inanimate object. The origins of the e ...
s surrounding
Polaris Polaris is a star in the northern circumpolar constellation of Ursa Minor. It is designated α Ursae Minoris ( Latinized to ''Alpha Ursae Minoris'') and is commonly called the North Star or Pole Star. With an apparent magnitude that ...
. These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.


Communication

Birds
communicate Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inquir ...
primarily using visual and auditory signals. Signals can be interspecific (between species) and intraspecific (within species). Birds sometimes use plumage to assess and assert social dominance, to display breeding condition in sexually selected species, or to make threatening displays, as in the
sunbittern The sunbittern (''Eurypyga helias'') is a bittern-like bird of tropical regions of the Americas, and the sole member of the family Eurypygidae (sometimes spelled Eurypigidae) and genus ''Eurypyga''. It is found in Central and South America, and ...
's mimicry of a large predator to ward off
hawk Hawks are bird of prey, birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica. * The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. Th ...
s and protect young chicks. Visual communication among birds may also involve ritualised displays, which have developed from non-signalling actions such as preening, the adjustments of feather position, pecking, or other behaviour. These displays may signal aggression or submission or may contribute to the formation of pair-bonds. The most elaborate displays occur during courtship, where "dances" are often formed from complex combinations of many possible component movements; males' breeding success may depend on the quality of such displays. Bird calls and songs, which are produced in the
syrinx In classical Greek mythology, Syrinx (Greek Σύριγξ) was a nymph and a follower of Artemis, known for her chastity. Pursued by the amorous god Pan, she ran to a river's edge and asked for assistance from the river nymphs. In answer, sh ...
, are the major means by which birds communicate with
sound In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' by the ...
. This communication can be very complex; some species can operate the two sides of the syrinx independently, allowing the simultaneous production of two different songs. Calls are used for a variety of purposes, including mate attraction, evaluation of potential mates, bond formation, the claiming and maintenance of territories, the identification of other individuals (such as when parents look for chicks in colonies or when mates reunite at the start of breeding season), and the warning of other birds of potential predators, sometimes with specific information about the nature of the threat. Some birds also use mechanical sounds for auditory communication. The ''
Coenocorypha The austral snipes, also known as the New Zealand snipes or tutukiwi, are a genus, ''Coenocorypha'', of tiny birds in the sandpiper family (biology), family, which are now only found on New Zealand outlying islands, New Zealand's outlying islands ...
''
snipe A snipe is any of about 26 wading bird species in three genera in the family Scolopacidae. They are characterized by a very long, slender bill, eyes placed high on the head, and cryptic/camouflaging plumage. The ''Gallinago'' snipes have a near ...
s of
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
drive air through their feathers,
woodpecker Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...
s drum for long-distance communication, and
palm cockatoo The palm cockatoo (''Probosciger aterrimus''), also known as the goliath cockatoo or great black cockatoo, is a large smoky-grey or black parrot of the cockatoo family native to New Guinea, Aru Islands, and Cape York Peninsula. It has a very l ...
s use tools to drum.


Flocking and other associations

While some birds are essentially territorial or live in small family groups, other birds may form large
flocks Flocking is the behaviour exhibited when a group of birds, called a flock, are foraging or in flight. Computer simulations and mathematical models that have been developed to emulate the flocking behaviours of birds can also generally be applie ...
. The principal benefits of flocking are
safety in numbers Safety in numbers is the hypothesis that, by being part of a large physical group or mass, an individual is less likely to be the victim of a mishap, accident, attack, or other bad event. Some related theories also argue (and can show statistica ...
and increased foraging efficiency. Defence against predators is particularly important in closed habitats like forests, where
ambush predation Ambush predators or sit-and-wait predators are carnivorous animals that capture or trap prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey us ...
is common and multiple eyes can provide a valuable
early warning system An early warning system is a warning system that can be implemented as a chain of information communication systems and comprises sensors, event detection and decision subsystems for early identification of hazards. They work together to for ...
. This has led to the development of many
mixed-species feeding flock A mixed-species feeding flock, also termed a mixed-species foraging flock, mixed hunting party or informally bird wave, is a flock of usually insectivorous birds of different species that join each other and move together while foraging. These ar ...
s, which are usually composed of small numbers of many species; these flocks provide safety in numbers but increase potential competition for resources. Costs of flocking include bullying of socially subordinate birds by more dominant birds and the reduction of feeding efficiency in certain cases. Birds sometimes also form associations with non-avian species. Plunge-diving
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s associate with
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the ...
s and
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length: ...
, which push shoaling fish towards the surface. Some species of
hornbill Hornbills (Bucerotidae) are a family (biology), family of bird found in tropical and subtropical Africa, Asia and Melanesia. They are characterized by a long, down-curved bill which is frequently brightly coloured and sometimes has a Casque (an ...
s have a mutualistic relationship with
dwarf mongoose The common dwarf mongoose (''Helogale parvula'') is a mongoose species native to Angola, northern Namibia, KwaZulu-Natal in South Africa, Zambia and East Africa. It is part of the genus ''Helogale'', along with the Ethiopian dwarf mongoose. Char ...
s, in which they forage together and warn each other of nearby
birds of prey Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predators ...
and other predators.


Resting and roosting

The high metabolic rates of birds during the active part of the day is supplemented by rest at other times. Sleeping birds often use a type of sleep known as vigilant sleep, where periods of rest are interspersed with quick eye-opening "peeks", allowing them to be sensitive to disturbances and enable rapid escape from threats.
Swift Swift or SWIFT most commonly refers to: * SWIFT, an international organization facilitating transactions between banks ** SWIFT code * Swift (programming language) * Swift (bird), a family of birds It may also refer to: Organizations * SWIFT, ...
s are believed to be able to sleep in flight and radar observations suggest that they orient themselves to face the wind in their roosting flight. It has been suggested that there may be certain kinds of sleep which are possible even when in flight. Some birds have also demonstrated the capacity to fall into
slow-wave sleep Slow-wave sleep (SWS), often referred to as deep sleep, consists of stage three of non-rapid eye movement sleep. It usually lasts between 70 and 90 minutes and takes place during the first hours of the night. Initially, SWS consisted of both St ...
one
hemisphere Hemisphere refers to: * A half of a sphere As half of the Earth * A hemisphere of Earth ** Northern Hemisphere ** Southern Hemisphere ** Eastern Hemisphere ** Western Hemisphere ** Land and water hemispheres * A half of the (geocentric) celes ...
of the brain at a time. The birds tend to exercise this ability depending upon its position relative to the outside of the flock. This may allow the eye opposite the sleeping hemisphere to remain vigilant for
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s by viewing the outer margins of the flock. This adaptation is also known from
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s.
Communal roosting Communal roosting is an animal behavior where a group of individuals, typically of the same species, congregate in an area for a few hours based on an external signal and will return to the same site with the reappearance of the signal. Environment ...
is common because it lowers the loss of body heat and decreases the risks associated with predators. Roosting sites are often chosen with regard to thermoregulation and safety. Unusual mobile roost sites include large herbivores on the African savanna that are used by
oxpecker The oxpeckers are two species of bird which make up the genus ''Buphagus'', and family Buphagidae. The oxpeckers were formerly usually treated as a subfamily, Buphaginae, within the starling family, Sturnidae, but molecular phylogenetic studi ...
s. Many sleeping birds bend their heads over their backs and tuck their bills in their back feathers, although others place their beaks among their breast feathers. Many birds rest on one leg, while some may pull up their legs into their feathers, especially in cold weather.
Perching birds A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
have a tendon-locking mechanism that helps them hold on to the perch when they are asleep. Many ground birds, such as quails and pheasants, roost in trees. A few parrots of the genus '' Loriculus'' roost hanging upside down. Some
hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s go into a nightly state of
torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature and metabolic rate. Torpor enables animals to survive periods of reduced food availability. The term "torpor" can refer to the time ...
accompanied with a reduction of their metabolic rates. This physiological adaptation shows in nearly a hundred other species, including
owlet-nightjar Owlet-nightjars are small crepuscular birds related to the nightjars and frogmouths. Most are native to New Guinea, but some species extend to Australia, the Moluccas, and New Caledonia. A flightless species from New Zealand is extinct. There is ...
s,
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called goatsuckers, due to the ancient folk ta ...
s, and
woodswallow Woodswallows are soft-plumaged, somber-coloured passerine birds in the genus ''Artamus''. The woodswallows are either treated as a subfamily, Artaminae, in an expanded family Artamidae (also including the subfamily Cracticinae), or as the only ge ...
s. One species, the
common poorwill The common poorwill (''Phalaenoptilus nuttallii'') is a nocturnal bird of the family Caprimulgidae, the nightjars. It is found from British Columbia and southeastern Alberta, through the western United States to northern Mexico. The bird's habi ...
, even enters a state of
hibernation Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
. Birds do not have sweat glands, but can lose water directly through the skin, and they may cool themselves by moving to shade, standing in water, panting, increasing their surface area, fluttering their throat or using special behaviours like
urohidrosis Urohidrosis (sometimes misspelled "urohydrosis") is the habit in some birds of defecating onto the scaly portions of the legs as a cooling mechanism, using evaporative cooling of the fluids. Several species of storks and New World vultures exhibi ...
to cool themselves.


Breeding


Social systems

Ninety-five per cent of bird species are socially monogamous. These species pair for at least the length of the breeding season or—in some cases—for several years or until the death of one mate. Monogamy allows for both
paternal care In biology, paternal care is parental investment provided by a male to his own offspring. It is a complex social behaviour in vertebrates associated with animal mating systems, life history traits, and ecology. Paternal care may be provided in co ...
and biparental care, which is especially important for species in which care from both the female and the male parent is required in order to successfully rear a brood. Among many socially monogamous species,
extra-pair copulation Extra-pair copulation (EPC) is a mating behaviour in monogamous species. Monogamy is the practice of having only one sexual partner at any one time, forming a long-term bond and combining efforts to raise offspring together; mating outside this pai ...
(infidelity) is common. Such behaviour typically occurs between dominant males and females paired with subordinate males, but may also be the result of
forced copulation Sexual coercion among animals is the use of violence, threats, harassment, and other tactics to help them forcefully copulate.Smuts, Barbara BMale Aggression and Sexual Coercion of Females in Nonhuman Primates and Other Mammals: Evidence and Theo ...
in ducks and other anatids. For females, possible benefits of extra-pair copulation include getting better genes for her offspring and insuring against the possibility of infertility in her mate. Males of species that engage in extra-pair copulations will closely guard their mates to ensure the parentage of the offspring that they raise. Other mating systems, including
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any o ...
,
polyandry Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
,
polygamy Crimes Polygamy (from Late Greek (') "state of marriage to many spouses") is the practice of marrying multiple spouses. When a man is married to more than one wife at the same time, sociologists call this polygyny. When a woman is married ...
,
polygynandry Polygynandry is a mating system in which both males and females have multiple mating partners during a breeding season. In sexually reproducing diploid animals, different mating strategies are employed by males and females, because the cost of game ...
, and
promiscuity Promiscuity is the practice of engaging in sexual activity frequently with different Sexual partner, partners or being indiscriminate in the choice of sexual partners. The term can carry a moral judgment. A common example of behavior viewed as pro ...
, also occur. Polygamous breeding systems arise when females are able to raise broods without the help of males. Mating systems vary across bird families but variations within species are thought to be driven by environmental conditions. Breeding usually involves some form of courtship display, typically performed by the male. Most displays are rather simple and involve some type of
song A song is a musical composition intended to be performed by the human voice. This is often done at distinct and fixed pitches (melodies) using patterns of sound and silence. Songs contain various forms, such as those including the repetitio ...
. Some displays, however, are quite elaborate. Depending on the species, these may include wing or tail drumming, dancing, aerial flights, or communal lekking. Females are generally the ones that drive partner selection, although in the polyandrous
phalaropes __NOTOC__ A phalarope is any of three living species of slender-necked shorebirds in the genus ''Phalaropus'' of the bird family Scolopacidae. Phalaropes are close relatives of the shanks and tattlers, the '' Actitis'' and Terek sandpipers, ...
, this is reversed: plainer males choose brightly coloured females.
Courtship feeding A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices. Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simp ...
, billing and are commonly performed between partners, generally after the birds have paired and mated. Homosexual behaviour has been observed in males or females in numerous species of birds, including copulation, pair-bonding, and joint parenting of chicks. Over 130 avian species around the world engage in sexual interactions between the same sex or homosexual behaviours. "Same-sex courtship activities may involve elaborate displays, synchronized dances, gift-giving ceremonies, or behaviors at specific display areas including bowers, arenas, or leks."


Territories, nesting and incubation

Many birds actively defend a territory from others of the same species during the breeding season; maintenance of territories protects the food source for their chicks. Species that are unable to defend feeding territories, such as
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s and
swift Swift or SWIFT most commonly refers to: * SWIFT, an international organization facilitating transactions between banks ** SWIFT code * Swift (programming language) * Swift (bird), a family of birds It may also refer to: Organizations * SWIFT, ...
s, often breed in
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
instead; this is thought to offer protection from predators. Colonial breeders defend small nesting sites, and competition between and within species for nesting sites can be intense. All birds lay
amniotic egg Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are distingu ...
s with hard shells made mostly of calcium carbonate. Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
d eggs. There are many exceptions to this pattern, however; the ground-nesting
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called goatsuckers, due to the ancient folk ta ...
s have pale eggs, and camouflage is instead provided by their plumage. Species that are victims of
brood parasites Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own ...
have varying egg colours to improve the chances of spotting a parasite's egg, which forces female parasites to match their eggs to those of their hosts. Bird eggs are usually laid in a
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materia ...
. Most species create somewhat elaborate nests, which can be cups, domes, plates, mounds, or burrows. Some bird nests can be a simple scrape, with minimal or no lining; most seabird and wader nests are no more than a scrape on the ground. Most birds build nests in sheltered, hidden areas to avoid predation, but large or colonial birds—which are more capable of defence—may build more open nests. During nest construction, some species seek out plant matter from plants with parasite-reducing toxins to improve chick survival, and feathers are often used for nest insulation. Some bird species have no nests; the cliff-nesting
common guillemot The common murre or common guillemot (''Uria aalge'') is a large auk. It has a circumpolar distribution, occurring in low-Arctic and boreal waters in the North Atlantic and North Pacific. It spends most of its time at sea, only coming to land to ...
lays its eggs on bare rock, and male
emperor penguin The emperor penguin (''Aptenodytes forsteri'') is the tallest and heaviest of all living penguin species and is endemic to Antarctica. The male and female are similar in plumage and size, reaching in length and weighing from . Feathers of th ...
s keep eggs between their body and feet. The absence of nests is especially prevalent in open habitat ground-nesting species where any addition of nest material would make the nest more conspicuous. Many ground nesting birds lay a clutch of eggs that hatch synchronously, with
precocial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
chicks led away from the nests ( nidifugous) by their parents soon after hatching. Incubation, which regulates temperature for chick development, usually begins after the last egg has been laid. In monogamous species incubation duties are often shared, whereas in polygamous species one parent is wholly responsible for incubation. Warmth from parents passes to the eggs through brood patches, areas of bare skin on the abdomen or breast of the incubating birds. Incubation can be an energetically demanding process; adult albatrosses, for instance, lose as much as of body weight per day of incubation. The warmth for the incubation of the eggs of
megapode The megapodes, also known as incubator birds or mound-builders, are stocky, medium-large, chicken-like birds with small heads and large feet in the family Megapodiidae. Their name literally means "large foot" and is a reference to the heavy legs ...
s comes from the sun, decaying vegetation or volcanic sources. Incubation periods range from 10 days (in
woodpecker Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...
s,
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
s and
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by t ...
birds) to over 80 days (in albatrosses and
kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...
s). The diversity of characteristics of birds is great, sometimes even in closely related species. Several avian characteristics are compared in the table below.


Parental care and fledging

At the time of their hatching, chicks range in development from helpless to independent, depending on their species. Helpless chicks are termed ''
altricial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
'', and tend to be born small, blind, immobile and naked; chicks that are mobile and feathered upon hatching are termed ''
precocial In biology, altricial species are those in which the young are underdeveloped at the time of birth, but with the aid of their parents mature after birth. Precocial species are those in which the young are relatively mature and mobile from the mome ...
''. Altricial chicks need help thermoregulating and must be brooded for longer than precocial chicks. The young of many bird species do not precisely fit into either the precocial or altricial category, having some aspects of each and thus fall somewhere on an "altricial-precocial spectrum". Chicks at neither extreme but favouring one or the other may be termed or . The length and nature of parental care varies widely amongst different orders and species. At one extreme, parental care in
megapode The megapodes, also known as incubator birds or mound-builders, are stocky, medium-large, chicken-like birds with small heads and large feet in the family Megapodiidae. Their name literally means "large foot" and is a reference to the heavy legs ...
s ends at hatching; the newly hatched chick digs itself out of the nest mound without parental assistance and can fend for itself immediately. At the other extreme, many seabirds have extended periods of parental care, the longest being that of the
great frigatebird The great frigatebird (''Fregata minor'') is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific (including the Galapagos Islands) and Indian Oceans, as well as a tiny population in the South At ...
, whose chicks take up to six months to
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
and are fed by the parents for up to an additional 14 months. The ''chick guard stage'' describes the period of breeding during which one of the adult birds is permanently present at the nest after chicks have hatched. The main purpose of the guard stage is to aid offspring to thermoregulate and protect them from predation.In some species, both parents care for nestlings and fledglings; in others, such care is the responsibility of only one sex. In some species,
other members Other often refers to: * Other (philosophy), a concept in psychology and philosophy Other or The Other may also refer to: Film and television * ''The Other'' (1913 film), a German silent film directed by Max Mack * ''The Other'' (1930 film), ...
of the same species—usually close relatives of the
breeding pair Breeding pair is a pair of animals which cooperate over time to produce offspring with some form of a bond between the individuals.Gaston, A. J.The evolution of group territorial behavior and cooperative breeding" The American Naturalist 112.988 ...
, such as offspring from previous broods—will help with the raising of the young. Such
alloparenting Alloparenting (also referred to as alloparental care) is a term used to classify any form of parental care provided by an individual towards young that aren't its own direct offspring. These are often referred to as "non-descendant" young, even th ...
is particularly common among the
Corvida The "Corvida" were one of two "parvorders" contained within the suborder Passeri, as proposed in the Sibley-Ahlquist taxonomy, the other being Passerida. Standard taxonomic practice would place them at the rank of infraorder. More recent resea ...
, which includes such birds as the true
crows The Common Remotely Operated Weapon Station (CROWS) is a series of remote weapon stations used by the US military on its armored vehicles and ships. It allows weapon operators to engage targets without leaving the protection of their vehicle. T ...
,
Australian magpie The Australian magpie (''Gymnorhina tibicen'') is a black and white passerine bird native to Australia and southern New Guinea. Although once considered to be three separate species, it is now considered to be one, with nine recognised subs ...
and
fairy-wren The Australasian wrens are a family, Maluridae, of small, insectivorous passerine birds endemic to Australia and New Guinea. While commonly known as wrens, they are unrelated to the true wrens. The family comprises 32 species (including sixteen ...
s, but has been observed in species as different as the
rifleman A rifleman is an infantry soldier armed with a rifling, rifled long gun. Although the rifleman role had its origin with 16th century hand cannoneers and 17th century musketeers, the term originated in the 18th century with the introduction o ...
and
red kite The red kite (''Milvus milvus'') is a medium-large bird of prey in the family Accipitridae, which also includes many other diurnal raptors such as eagles, buzzards, and harriers. The species currently breeds in the Western Palearctic region o ...
. Among most groups of animals, male parental care is rare. In birds, however, it is quite common—more so than in any other vertebrate class. Although territory and nest site defence, incubation, and chick feeding are often shared tasks, there is sometimes a
division of labour The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise (specialisation). Individuals, organizations, and nations are endowed with, or acquire specialised capabilities, and ...
in which one mate undertakes all or most of a particular duty. The point at which chicks
fledge Fledging is the stage in a flying animal's life between hatching or birth and becoming capable of flight. This term is most frequently applied to birds, but is also used for bats. For altricial birds, those that spend more time in vulnerable c ...
varies dramatically. The chicks of the '' Synthliboramphus'' murrelets, like the
ancient murrelet The ancient murrelet (') is a bird in the auk family. The English term "murrelet" is a diminutive of "murre", a word of uncertain origins, but which may imitate the call of the common guillemot. Ancient murrelets are called "ancient" because th ...
, leave the nest the night after they hatch, following their parents out to sea, where they are raised away from terrestrial predators. Some other species, such as ducks, move their chicks away from the nest at an early age. In most species, chicks leave the nest just before, or soon after, they are able to fly. The amount of parental care after fledging varies; albatross chicks leave the nest on their own and receive no further help, while other species continue some supplementary feeding after fledging. Chicks may also follow their parents during their first
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
.


Brood parasites

Brood parasitism Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own ...
, in which an egg-layer leaves her eggs with another individual's brood, is more common among birds than any other type of organism. After a parasitic bird lays her eggs in another bird's nest, they are often accepted and raised by the host at the expense of the host's own brood. Brood parasites may be either ''obligate brood parasites'', which must lay their eggs in the nests of other species because they are incapable of raising their own young, or ''non-obligate brood parasites'', which sometimes lay eggs in the nests of
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...
s to increase their reproductive output even though they could have raised their own young. One hundred bird species, including
honeyguide Honeyguides (family Indicatoridae) are near passerine birds in the order Piciformes. They are also known as indicator birds, or honey birds, although the latter term is also used more narrowly to refer to species of the genus '' Prodotiscus''. Th ...
s, icterids, and
ducks Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
, are obligate parasites, though the most famous are the
cuckoo Cuckoos are birds in the Cuculidae family, the sole taxon in the order Cuculiformes . The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals and anis. The coucals and anis are sometimes separ ...
s. Some brood parasites are adapted to hatch before their host's young, which allows them to destroy the host's eggs by pushing them out of the nest or to kill the host's chicks; this ensures that all food brought to the nest will be fed to the parasitic chicks.


Sexual selection

Birds have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
a variety of
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
behaviours, with the
peacock Peafowl is a common name for three bird species in the genera ''Pavo (genus), Pavo'' and ''Afropavo'' within the tribe Pavonini of the family Phasianidae, the pheasants and their allies. Male peafowl are referred to as peacocks, and female pea ...
tail being perhaps the most famous example of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
and the
Fisherian runaway Fisherian runaway or runaway selection is a sexual selection mechanism proposed by the mathematical biologist Ronald Fisher in the early 20th century, to account for the evolution of ostentatious male ornamentation by persistent, directional fe ...
. Commonly occurring
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
s such as size and colour differences are energetically costly attributes that signal competitive breeding situations. Many types of avian
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...
have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signalling adaptations ensure that sexually selected ornaments such as plumage colouration and courtship behaviour are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviours.


Inbreeding depression

Inbreeding causes early death (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
) in the
zebra finch The zebra finches are two species of estrildid finch in the genus ''Taeniopygia'' found in Australia and Indonesia. They are seed-eaters that travel in large flocks. The species are: Previously, both species were classified as a single species ...
''Taeniopygia guttata''. Embryo survival (that is, hatching success of fertile eggs) was significantly lower for sib-sib mating pairs than for unrelated pairs.
Darwin's finch Darwin's finches (also known as the Galápagos finches) are a group of about 18 species of passerine birds. They are well known for their remarkable diversity in beak form and function. They are often classified as the subfamily Geospizinae or t ...
''Geospiza scandens'' experiences
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
(reduced survival of offspring) and the magnitude of this effect is influenced by environmental conditions such as low food availability.


Inbreeding avoidance

Incestuous matings by the purple-crowned fairy wren ''Malurus coronatus'' result in severe fitness costs due to
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
(greater than 30% reduction in hatchability of eggs). Females paired with related males may undertake extra pair matings (see Promiscuity#Other animals for 90% frequency in avian species) that can reduce the negative effects of inbreeding. However, there are ecological and demographic constraints on extra pair matings. Nevertheless, 43% of broods produced by incestuously paired females contained extra pair young. Inbreeding depression occurs in the
great tit The great tit (''Parus major'') is a passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central Asia and east across the Palearctic to the Amur River, south to parts of North Af ...
(''Parus major'') when the offspring produced as a result of a mating between close relatives show reduced fitness. In natural populations of ''Parus major'', inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative.
Southern pied babbler The southern pied babbler (''Turdoides bicolor'') is a species of bird in the family Leiothrichidae, found in dry savannah of Botswana, Namibia, South Africa, and Zimbabwe. Description The southern pied babbler is a medium-sized cooperatively b ...
s ''Turdoides bicolor'' appear to avoid inbreeding in two ways. The first is through dispersal, and the second is by avoiding familiar group members as mates.
Cooperative breeding Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers. Cooperative breeding encompasses a wide variety of group ...
in birds typically occurs when offspring, usually males, delay dispersal from their natal group in order to remain with the family to help rear younger kin. Female offspring rarely stay at home, dispersing over distances that allow them to breed independently, or to join unrelated groups. In general, inbreeding is avoided because it leads to a reduction in progeny fitness (
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
) due largely to the homozygous expression of deleterious recessive alleles.
Cross-fertilisation Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...
between unrelated individuals ordinarily leads to the masking of deleterious recessive alleles in progeny.


Ecology

Birds occupy a wide range of ecological positions. While some birds are generalists, others are highly specialised in their habitat or food requirements. Even within a single habitat, such as a forest, the niches occupied by different species of birds vary, with some species feeding in the
forest canopy In biology, the canopy is the aboveground portion of a plant cropping or crop, formed by the collection of individual plant crowns. In forest ecology, canopy also refers to the upper layer or habitat zone, formed by mature tree crowns an ...
, others beneath the canopy, and still others on the forest floor. Forest birds may be
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
s,
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance an ...
s, or
nectarivore In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants. Nectar as a food source presents a number of benefits a ...
s. Aquatic birds generally feed by fishing, plant eating, and piracy or
kleptoparasitism Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when foo ...
. Many grassland birds are granivores. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s. Birds are also preyed upon by a range of mammals including a few avivorous bats. A wide range of endo- and ectoparasites depend on birds and some parasites that are transmitted from parent to young have co-evolved and show host-specificity. Some nectar-feeding birds are important pollinators, and many frugivores play a key role in seed dispersal. Plants and pollinating birds often coevolve, and in some cases a flower's primary pollinator is the only species capable of reaching its nectar. Birds are often important to island ecology. Birds have frequently reached islands that mammals have not; on those islands, birds may fulfil ecological roles typically played by larger animals. For example, in New Zealand nine species of
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
were important browsers, as are the
kererū The kererū (''Hemiphaga novaeseelandiae'') or New Zealand pigeon is a species of pigeon native to New Zealand. Johann Friedrich Gmelin described the bird in 1789 as a large, conspicuous pigeon up to in length and in weight, with a white br ...
and kokako today. Today the plants of New Zealand retain the defensive adaptations evolved to protect them from the extinct moa. Many birds act as
ecosystem engineer An ecosystem engineer is any species that creates, significantly modifies, maintains or destroys a habitat. These organisms can have a large impact on species richness and landscape-level heterogeneity of an area. As a result, ecosystem enginee ...
s through the construction of nests, which provide important microhabitats and food for hundreds of species of invertebrates. Nesting
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s may affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
, which may enrich the local soil and the surrounding seas. A wide variety of
avian ecology field methods There are many field methods available for conducting avian ecological research. They can be divided into three types: counts, nest monitoring, and capturing and marking. Basic counts Basic bird counts are a good way to estimate population size, ...
, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology.


Relationship with humans

Since birds are highly visible and common animals, humans have had a relationship with them since the dawn of man. Sometimes, these relationships are mutualistic, like the cooperative honey-gathering among
honeyguide Honeyguides (family Indicatoridae) are near passerine birds in the order Piciformes. They are also known as indicator birds, or honey birds, although the latter term is also used more narrowly to refer to species of the genus '' Prodotiscus''. Th ...
s and African peoples such as the Borana. Other times, they may be
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit fro ...
, as when species such as the
house sparrow The house sparrow (''Passer domesticus'') is a bird of the sparrow family Passeridae, found in most parts of the world. It is a small bird that has a typical length of and a mass of . Females and young birds are coloured pale brown and grey, a ...
have benefited from human activities. Several bird species have become commercially significant agricultural pests, and some pose an aviation hazard. Human activities can also be detrimental, and have threatened numerous bird species with extinction (
hunting Hunting is the human activity, human practice of seeking, pursuing, capturing, or killing wildlife or feral animals. The most common reasons for humans to hunt are to harvest food (i.e. meat) and useful animal products (fur/hide (skin), hide, ...
,
avian lead poisoning Animal lead poisoning (also known as avian plumbism, or avian saturnism for birds) is a veterinary condition and pathology caused by increased levels of the heavy metal lead in animal's body. Lead interferes with a variety of body and natural proc ...
,
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
s,
roadkill Roadkill is an animal or animals that have been struck and killed by drivers of motor vehicles on highways. Wildlife-vehicle collisions (WVC) have increasingly been the topic of academic research to understand the causes, and how it can be mi ...
,
wind turbine A wind turbine is a device that converts the kinetic energy of wind into electrical energy. Hundreds of thousands of large turbines, in installations known as wind farms, now generate over 650 gigawatts of power, with 60 GW added each year. ...
kills and predation by pet
cat The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of ...
s and
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
s are common causes of death for birds). Birds can act as vectors for spreading diseases such as psittacosis,
salmonellosis Salmonellosis, more commonly known as food poisoning is a symptomatic infection caused by bacteria of the ''Salmonella'' type. It is also a food-borne disease and are defined as diseases, usually either infectious or toxic in nature, caused by a ...
,
campylobacteriosis Campylobacteriosis is an infection by the '' Campylobacter'' bacterium, most commonly ''C. jejuni''. It is among the most common bacterial infections of humans, often a foodborne illness. It produces an inflammatory, sometimes bloody, diarrhea or ...
, mycobacteriosis (avian
tuberculosis Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, in ...
),
avian influenza Avian influenza, known informally as avian flu or bird flu, is a variety of influenza caused by viruses adapted to birds.
(bird flu),
giardiasis Giardiasis is a parasitic disease caused by ''Giardia duodenalis'' (also known as ''G. lamblia'' and ''G. intestinalis''). Infected individuals who experience symptoms (about 10% have no symptoms) may have diarrhea, abdominal pain, and weight los ...
, and
cryptosporidiosis Cryptosporidiosis, sometimes informally called crypto, is a parasitic disease caused by ''Cryptosporidium'', a genus of protozoan parasites in the phylum Apicomplexa. It affects the distal small intestine and can affect the respiratory tract in ...
over long distances. Some of these are
zoonotic diseases A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a bacterium, virus, parasite or prion) that has jumped from a non-human (usually a vertebrate) to a human. ...
that can also be transmitted to humans.


Economic importance

Domesticated birds raised for meat and eggs, called
poultry Poultry () are domesticated birds kept by humans for their eggs, their meat or their feathers. These birds are most typically members of the superorder Galloanserae (fowl), especially the order Galliformes (which includes chickens, quails, a ...
, are the largest source of animal protein eaten by humans; in 2003, tons of poultry and tons of eggs were produced worldwide.
Chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adult m ...
s account for much of human poultry consumption, though domesticated
turkeys The turkey is a large bird in the genus ''Meleagris'', native to North America. There are two extant turkey species: the wild turkey (''Meleagris gallopavo'') of eastern and central North America and the ocellated turkey (''Meleagris ocellat ...
,
ducks Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
, and
geese A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...
are also relatively common. Many species of birds are also hunted for meat. Bird hunting is primarily a recreational activity except in extremely undeveloped areas. The most important birds hunted in North and South America are waterfowl; other widely hunted birds include
pheasant Pheasants ( ) are birds of several genera within the family (biology), family Phasianidae in the order (biology), order Galliformes. Although they can be found all over the world in introduced (and captive) populations, the pheasant genera na ...
s,
wild turkey The wild turkey (''Meleagris gallopavo'') is an Upland game bird, upland ground bird native to North America, one of two extant species of Turkey (bird), turkey and the heaviest member of the order Galliformes. It is the ancestor to the domestic ...
s, quail,
dove Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
s,
partridge A partridge is a medium-sized Galliformes, galliform bird in any of several genera, with a wide Indigenous (ecology), native distribution throughout parts of Europe, Asia and Africa. Several species have been introduced to the Americas. They a ...
,
grouse Grouse are a group of birds from the order Galliformes, in the family Phasianidae. Grouse are presently assigned to the tribe Tetraonini (formerly the subfamily Tetraoninae and the family Tetraonidae), a classification supported by mitochondr ...
,
snipe A snipe is any of about 26 wading bird species in three genera in the family Scolopacidae. They are characterized by a very long, slender bill, eyes placed high on the head, and cryptic/camouflaging plumage. The ''Gallinago'' snipes have a near ...
, and
woodcock The woodcocks are a group of seven or eight very similar living species of wading birds in the genus ''Scolopax''. The genus name is Latin for a snipe or woodcock, and until around 1800 was used to refer to a variety of waders. The English name ...
.
Muttonbirding Muttonbirding is the seasonal harvesting of the chicks of petrels, especially shearwater species, for food, oil and feathers by recreational or commercial hunters. Such hunting of petrels and other seabirds has occurred in various locations since p ...
is also popular in Australia and New Zealand. Although some hunting, such as that of muttonbirds, may be sustainable, hunting has led to the extinction or endangerment of dozens of species. Other commercially valuable products from birds include feathers (especially the down of geese and ducks), which are used as insulation in clothing and bedding, and seabird faeces (
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
), which is a valuable source of phosphorus and nitrogen. The
War of the Pacific The War of the Pacific ( es, link=no, Guerra del Pacífico), also known as the Saltpeter War ( es, link=no, Guerra del salitre) and by multiple other names, was a war between Chile and a Bolivian–Peruvian alliance from 1879 to 1884. Fought ...
, sometimes called the Guano War, was fought in part over the control of guano deposits. Birds have been domesticated by humans both as pets and for practical purposes. Colourful birds, such as
parrots Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genus (biology), genera comprising the order (biology), order Psittaciformes (), found mostly in tropics, tropical and subtropics, subtropical regions. The order ...
and
myna The myna (; also spelled mynah) is a bird of the starling family (Sturnidae). This is a group of passerine birds which are native to southern Asia, especially India, Pakistan and Bangladesh. Several species have been introduced to areas like ...
s, are bred in
captivity Captivity, or being held captive, is a state wherein humans or other animals are confined to a particular space and prevented from leaving or moving freely. An example in humans is imprisonment. Prisoners of war are usually held in captivity by a ...
or kept as pets, a practice that has led to the illegal trafficking of some
endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
.
Falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Falcons are widely distributed on all continents of the world except Antarctica, though closely related raptors did occur there in the Eocene. Adult falcons ...
s and
cormorant Phalacrocoracidae is a family of approximately 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the IOC adopted a consensus taxonomy of seven ge ...
s have long been used for
hunting Hunting is the human activity, human practice of seeking, pursuing, capturing, or killing wildlife or feral animals. The most common reasons for humans to hunt are to harvest food (i.e. meat) and useful animal products (fur/hide (skin), hide, ...
and
fishing Fishing is the activity of trying to catch fish. Fish are often caught as wildlife from the natural environment, but may also be caught from stocked bodies of water such as ponds, canals, park wetlands and reservoirs. Fishing techniques inclu ...
, respectively.
Messenger pigeon The homing pigeon, also called the mail pigeon or messenger pigeon, is a variety of domestic pigeons (''Columba livia domestica'') derived from the wild rock dove, selectively bred for its ability to find its way home over extremely long distan ...
s, used since at least 1 AD, remained important as recently as
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...
. Today, such activities are more common either as hobbies, for entertainment and tourism, Amateur bird enthusiasts (called birdwatchers, twitchers or, more commonly,
birders Birdwatching, or birding, is the observing of birds, either as a recreational activity or as a form of citizen science. A birdwatcher may observe by using their naked eye, by using a visual enhancement device like binoculars or a telescope, by ...
) number in the millions. Many homeowners erect
bird feeder A birdfeeder, bird table, or tray feeder are devices placed outdoors to supply bird food to birds (bird feeding). The success of a bird feeder in attracting birds depends upon its placement and the kinds of foods offered, as different species h ...
s near their homes to attract various species.
Bird feeding Bird feeding is the activity of feeding wild birds, often by means of bird feeders. With a recorded history dating to the 6th century, the feeding of wild birds has been encouraged and celebrated in the United States and United Kingdom, with it b ...
has grown into a multimillion-dollar industry; for example, an estimated 75% of households in Britain provide food for birds at some point during the winter.


In religion and mythology

Birds play prominent and diverse roles in religion and mythology. In religion, birds may serve as either messengers or priests and leaders for a
deity A deity or god is a supernatural being who is considered divine or sacred. The ''Oxford Dictionary of English'' defines deity as a god or goddess, or anything revered as divine. C. Scott Littleton defines a deity as "a being with powers greate ...
, such as in the Cult of
Makemake Makemake (minor-planet designation 136472 Makemake) is a dwarf planet and – depending on how they are defined – the second-largest Kuiper belt object in the classical population, with a diameter approximately 60% that of Pluto. It ...
, in which the
Tangata manu The ''Tangata manu'' ("bird-man," from "human beings" + "bird") was the winner of a traditional competition on Rapa Nui (Easter Island). The ritual was an annual competition to collect the first sooty tern () egg of the season from the islet of ...
of
Easter Island Easter Island ( rap, Rapa Nui; es, Isla de Pascua) is an island and special territory of Chile in the southeastern Pacific Ocean, at the southeasternmost point of the Polynesian Triangle in Oceania. The island is most famous for its nearl ...
served as chiefs or as attendants, as in the case of
Hugin and Munin In Norse mythology, Huginn (Old Norse: "thought"Orchard (1997:92).) and Muninn (Old Norse "memory"Orchard (1997:115). or "mind"Lindow (2001:186).) are a pair of ravens that fly all over the world, Midgard, and bring information to the god Odin. ...
, the two
common raven The common raven (''Corvus corax'') is a large all-black passerine bird. It is the most widely distributed of all corvids, found across the Northern Hemisphere. It is a raven known by many names at the subspecies level; there are at least e ...
s who whispered news into the ears of the
Norse god In Germanic paganism, the indigenous religion of the ancient Germanic peoples who inhabited Germanic Europe, there were a number of different gods and goddesses. Germanic deities are attested from numerous sources, including works of literature, ...
Odin Odin (; from non, Óðinn, ) is a widely revered Æsir, god in Germanic paganism. Norse mythology, the source of most surviving information about him, associates him with wisdom, healing, death, royalty, the gallows, knowledge, war, battle, v ...
. In several civilisations of
ancient Italy The history of Italy covers the ancient period, the Middle Ages, and the modern era. Since classical antiquity, ancient Etruscans, various Italic peoples (such as the Latins, Samnites, and Umbri), Celts, '' Magna Graecia'' colonists, and oth ...
, particularly
Etruscan __NOTOC__ Etruscan may refer to: Ancient civilization *The Etruscan language, an extinct language in ancient Italy *Something derived from or related to the Etruscan civilization **Etruscan architecture **Etruscan art **Etruscan cities ** Etrusca ...
and
Roman Roman or Romans most often refers to: *Rome, the capital city of Italy *Ancient Rome, Roman civilization from 8th century BC to 5th century AD *Roman people, the people of ancient Rome *''Epistle to the Romans'', shortened to ''Romans'', a letter ...
religion Religion is usually defined as a social- cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatural, ...
, priests were involved in
augur An augur was a priest and official in the classical Roman world. His main role was the practice of augury, the interpretation of the will of the gods by studying the flight of birds. Determinations were based upon whether they were flying i ...
y, or interpreting the words of birds while the "auspex" (from which the word "auspicious" is derived) watched their activities to foretell events. They may also serve as
religious symbols A religious symbol is an iconic representation intended to represent a specific religion, or a specific concept within a given religion. Religious symbols have been used in the military in many countries, such as the United States military chapl ...
, as when
Jonah Jonah or Jonas, ''Yōnā'', "dove"; gr, Ἰωνᾶς ''Iōnâs''; ar, يونس ' or '; Latin: ''Ionas'' son of Amittai, is a prophet in the Hebrew Bible and the Quran, from Gath-hepher of the northern kingdom of Israel in about the 8th cent ...
(,
dove Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
) embodied the fright, passivity, mourning, and beauty traditionally associated with doves. Birds have themselves been deified, as in the case of the
common peacock The Indian peafowl (''Pavo cristatus''), also known as the common peafowl, and blue peafowl, is a peafowl species native to the Indian subcontinent. It has been introduced to many other countries. Male peafowl are referred to as peacocks, and ...
, which is perceived as Mother Earth by the people of southern India. In the ancient world, doves were used as symbols of the
Mesopotamian goddess Deities in ancient Mesopotamia were almost exclusively anthropomorphic. They were thought to possess extraordinary powers and were often envisioned as being of tremendous physical size. The deities typically wore ''melam'', an ambiguous substan ...
Inanna Inanna, also sux, 𒀭𒊩𒌆𒀭𒈾, nin-an-na, label=none is an List of Mesopotamian deities, ancient Mesopotamian goddess of love, war, and fertility. She is also associated with beauty, sex, Divine law, divine justice, and political p ...
(later known as Ishtar), the Canaanite mother goddess
Asherah Asherah (; he, אֲשֵׁרָה, translit=Ăšērā; uga, 𐎀𐎘𐎗𐎚, translit=ʾAṯiratu; akk, 𒀀𒅆𒋥, translit=Aširat; Qatabanian language, Qatabanian: ') in ancient Semitic religion, is a fertility goddess who appears in a ...
, and the Greek goddess
Aphrodite Aphrodite ( ; grc-gre, Ἀφροδίτη, Aphrodítē; , , ) is an ancient Greek goddess associated with love, lust, beauty, pleasure, passion, and procreation. She was syncretized with the Roman goddess . Aphrodite's major symbols include ...
. In
ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
,
Athena Athena or Athene, often given the epithet Pallas, is an ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretized with the Roman goddess Minerva. Athena was regarded as the patron and protectress of ...
, the goddess of wisdom and patron deity of the city of
Athens Athens ( ; el, Αθήνα, Athína ; grc, Ἀθῆναι, Athênai (pl.) ) is both the capital and largest city of Greece. With a population close to four million, it is also the seventh largest city in the European Union. Athens dominates ...
, had a
little owl The little owl (''Athene noctua''), also known as the owl of Athena or owl of Minerva, is a bird that inhabits much of the temperate and warmer parts of Europe, the Palearctic east to Korea, and North Africa. It was introduced into Britain at ...
as her symbol. In religious images preserved from the Inca and Tiwanaku empires, birds are depicted in the process of transgressing boundaries between earthly and underground spiritual realms. Indigenous peoples of the central Andes maintain legends of birds passing to and from metaphysical worlds.


In culture and folklore

Birds have featured in culture and art since prehistoric times, when they were represented in early
cave painting In archaeology, Cave paintings are a type of parietal art (which category also includes petroglyphs, or engravings), found on the wall or ceilings of caves. The term usually implies prehistoric origin, and the oldest known are more than 40,000 ye ...
s. Some birds have been perceived as monsters, including the mythological Roc and the
Māori Māori or Maori can refer to: Relating to the Māori people * Māori people of New Zealand, or members of that group * Māori language, the language of the Māori people of New Zealand * Māori culture * Cook Islanders, the Māori people of the C ...
's legendary , a giant bird capable of snatching humans. Birds were later used as symbols of power, as in the magnificent
Peacock Throne The Peacock Throne ( Hindustani: ''Mayūrāsana'', Sanskrit: मयूरासन, Urdu: تخت طاؤس, fa, تخت طاووس, ''Takht-i Tāvūs'') was a famous jewelled throne that was the seat of the emperors of the Mughal Empire in India ...
of the Mughal and
Persian Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...
emperors. With the advent of scientific interest in birds, many paintings of birds were commissioned for books. Among the most famous of these bird artists was
John James Audubon John James Audubon (born Jean-Jacques Rabin; April 26, 1785 – January 27, 1851) was an American self-trained artist, naturalist, and ornithologist. His combined interests in art and ornithology turned into a plan to make a complete pictoria ...
, whose paintings of North American birds were a great commercial success in Europe and who later lent his name to the
National Audubon Society The National Audubon Society (Audubon; ) is an American non-profit environmental organization dedicated to conservation of birds and their habitats. Located in the United States and incorporated in 1905, Audubon is one of the oldest of such organ ...
. Birds are also important figures in poetry; for example,
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
incorporated nightingales into his ''Odyssey'', and Catullus used a Old World sparrow, sparrow as an erotic symbol in his Catullus 2. The relationship between an albatross and a sailor is the central theme of Samuel Taylor Coleridge's ''The Rime of the Ancient Mariner'', which led to the use of the Albatross (metaphor), term as a metaphor for a 'burden'. Other English language, English metaphors derive from birds; vulture funds and vulture investors, for instance, take their name from the scavenging vulture. Perceptions of bird species vary across cultures. Owls are associated with bad luck, witchcraft, and death in parts of Africa, but are regarded as wise across much of Europe. Hoopoes were considered sacred in Ancient Egypt and symbols of virtue in Persia, but were thought of as thieves across much of Europe and harbingers of war in Scandinavia. In heraldry, birds, especially Eagle (heraldry), eagles, often appear in coats of arms.


In music

Birds in music, In music, birdsong has influenced composers and musicians in several ways: they can be inspired by birdsong; they can intentionally imitate bird song in a composition, as Antonio Vivaldi, Vivaldi, Olivier Messiaen, Messiaen, and Beethoven did, along with many later composers; they can incorporate recordings of birds into their works, as Ottorino Respighi first did; or like Beatrice Harrison and David Rothenberg, they can duet with birds.


Conservation

Although human activities have allowed the expansion of a few species, such as the barn swallow and European starling, they have caused population decreases or
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
in many other species. Over a hundred bird species have gone extinct in historical times, although the most dramatic human-caused avian extinctions, eradicating an estimated 750–1800 species, occurred during the human colonisation of Melanesian, Polynesian, and Micronesian islands. Many bird populations are declining worldwide, with 1,227 species listed as threatened species, threatened by BirdLife International and the IUCN in 2009. The most commonly cited human threat to birds is Habitat destruction, habitat loss. Other threats include overhunting, accidental mortality due to collisions with bird-skyscraper collisions, buildings or bird strike, vehicles, long-line fishing bycatch, pollution (including oil spills and pesticide use), competition and predation from nonnative invasive species, and Climate change and birds, climate change. Governments and conservation biology, conservation groups work to protect birds, either by passing laws that In-situ conservation, preserve and ecological restoration, restore bird habitat or by establishing Ex-situ conservation, captive populations for reintroductions. Such projects have produced some successes; one study estimated that conservation efforts saved 16 species of bird that would otherwise have gone extinct between 1994 and 2004, including the California condor and Norfolk parakeet.


See also

* Animal track * Avian sleep * Bat * Climate change and birds * Glossary of bird terms * List of individual birds * Ornithology * Paleocene dinosaurs


References


Further reading

* Roger Lederer und Carol Burr: ''Latein für Vogelbeobachter: über 3000 ornithologische Begriffe erklärt und erforscht'', aus dem Englischen übersetzt von Susanne Kuhlmannn-Krieg, Verlag DuMont, Köln 2014, . * del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.): ''Handbook of the Birds of the World'' (17-volume encyclopaedia), Lynx Edicions, Barcelona, 1992–2010. (''Vol. 1: Ostrich to Ducks'': , etc.). * ''All the Birds of the World'', Lynx Edicions, 2020. * ''National Geographic Field Guide to Birds of North America'', National Geographic, 7th edition, 2017. * ''National Audubon Society Field Guide to North American Birds: Eastern Region'', National Audubon Society, Knopf. * ''National Audubon Society Field Guide to North American Birds: Western Region'', National Audubon Society, Knopf. * Svensson, Lars: ''Birds of Europe'', Princeton University Press, second edition, 2010. * Svensson, Lars: ''Collins Bird Guide: The Most Complete Guide to the Birds of Britain and Europe'', Collins, 2nd edition, 2010.


External links


Birdlife International
– Dedicated to bird conservation worldwide; has a database with about 250,000 records on endangered bird species.



from the
National Audubon Society The National Audubon Society (Audubon; ) is an American non-profit environmental organization dedicated to conservation of birds and their habitats. Located in the United States and incorporated in 1905, Audubon is one of the oldest of such organ ...

Cornell Lab of Ornithology
*




Ornithology

Sora
– Searchable online research archive; Archives of the following ornithological journals ''The Auk'', ''Condor (journal), Condor'', ''Journal of Field Ornithology, ''North American Bird Bander'', ''Studies in Avian Biology'', ''Pacific Coast Avifauna'', and the ''the Wilson Bulletin, Wilson Bulletin''.
The Internet Bird Collection
– A free library of videos of the world's birds
The Institute for Bird Populations, California

List of field guides to birds
from the International Field Guides database
RSPB bird identifier
– Interactive identification of all UK birds

— University of California Museum of Paleontology. {{Authority control Birds, Animal classes Dinosaurs Extant Late Cretaceous first appearances Feathered dinosaurs Santonian first appearances Taxa named by Carl Linnaeus